"THE HEAD-CARIULE 
OF 


COLEOPTERR 
| 


Fenner Satterwaith Stickney 


THE HEAD-CAPSULE OF COLEOPTERA 


BY 


FENNER SATTERTHWAITE STICKNEY 
B. S. University of California, 1916 
M. S. University of Illinois, 1918 


THESIS 


Submitted in Partial Fulfillment of the Requirements for the 


Degree of 


DOCTOR OF PHILOSOPHY 


IN ENTOMOLOGY 


THE GRADUATE SCHOOL 
OF THE 


UNIVERSITY OF ILLINOIS 


1921 


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UNIVE RSATY OF ILLINOIS 


THE GRADUATE SCHOOL 


__May 14, 1921 19) _ 


I HEREBY RECOMMEND THAT THE THESIS PREPARED UNDER MY 


SUPERVISION BY.____Fenner Satterthwaite Stickney _ 


ENTITLED____ Tne Head-capsule of Coleoptera : 


BE ACCEPTED AS FULFILLING THIS PART OF THE REQUIREMENTS FOR 


THE DEGREE OF Doctor of Philosophy  —s_—> 


in ane of Thesis 


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Head of Department 


Recommendation concurred in* 


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I. Introduction 

II. Acknowledgments 

TIT, Materials 

IV. Head-capsule 
Epicranial suture 
Vertex 
Occipital suture 
Occiput 
Compound eyes 
Oculatea 
Supratentorina 
Ocelli 
Antennarisa 
Antacoria 
Antacava 


Pretentorina 


Postelypens 
Clypealia 
Mandibularia 
Preclypens 

Labrum 

Occipital foramen 


Submentum 


Metatentorina 


Maxilisria 


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Cervix 


Cervical sclerites 


EKpicranium 
Paracoila 
Postcoila 
Precoila 
Tentorium 
Pretentorium 
Metatentorium 
Corpotentorium 


Laminatentorium 


Bibliography 


Plates I = 4XVIT 


Vita 


Digitized by the Internet Archive 
in 2016 


httos://archive.org/details/neadcapsuleofcolOOstic 


I INTRODUCTION 
LeConte and Horn published in 1883 a classification of the Cole- 
optera that has stood the wear of time remarkably well. Since then, 
} however, a number of new classifications have been proposed: Lameere 
(1900 and 1903), Ganglbauer (1892-1904), Handlirsch (1906-1908), 
} Kolbe (1901,1908,and 1911), Sharp (1909), and Gahan (1911), all of 
| which differ more or less seriously in one way or another, and show, 


|'for one thing, the need of further comparative morphological data, 


/fi which te, of course, indispensable to the building of any thorough 


classification. lLeng's recent catalogue (1920) also emphasizes this 
|) need. 


A review of the literature seems to show but few studies based 


p : ; _ “of 
/}on the comparative morphology of a comprehensive series,coleopterous 


families. A number of European workers have published comparative 
studies of the wings of Coleoptera, the most recent being by d'Orchy- 
Hmont (1920). Sharp and Muir (1912) and Muir (1918) have published 

| the results of their investigations on the male genital tube in Cole- 
optera. Various internal structures have been discussed from time 

to time by a number of workers. Narrower in scope is the work of 
d'Orchymont (1916) on the classification of the Hydrophiloidea, based 
on a study of both the adult and the larva. Hyslop (1917), Béving 
and Champlain (1920), Craighead (1920), and Gage (1920), have pub- 
lished papers on the comparative morphology of various families, 
based on a study of the larvae. There are probably other comparative 
papers more or less extensive in scope, but I have not been able to 
find any such literature based on a study of the head-capsule. The 


comparative morphology of the head-capsule of some other orders, 


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however, has been investigated: Peterson (1915) on the Thysanoptera, 
Peterson (1916) on the Diptera, Yuasa (1920) on the Orthoptera, and 
| Hoke (1922) on the Plecoptera. These simply draw attention to the 
need of such an investigation of the head-capsule of Coleoptera, 
With the broader vision in mind of a more satisfactory and nat- 
ural classification of the Coleoptera, the following study on the 
l comparative morphology of the head-capsule is offered. This study 
does not aim by any means to exhaust the subject. There have been 
too few species investigated in each family to justify the making of 
any sweeping statements. This study can simply point out character- | 
istic conditions of structures as found in the different species of 


the families studied, revealing, therefore, inharmonies, and perhaps 


| suggesting improvments on the present arrangement of the classifica- 


tion. 

In order to reach a correct estimate of the degree of speciali- 
zation of the various parts of the head-capsule, an hypothetical 
type, representing a supposed primitive condition, has been construc 
ed. The structure of this hypothetical type is based on the struc- 
ture of the head-capsule of generalized insects and of generalized 
adult and larval Coleoptera. bach structure has been treated separ- 
ately, starting from the hypothetical type. The submentum has been 
ineluded in this study because of its bearing on certain develop- 
mental processes. All statements made refer to the species listed 
under "materials" only. The material studied was soaked in a 10% 
solution of potassium hydroxide until clarified, then washed in dis- 
tilled water to remove the hydroxide, and preserved in 70% alcohol. 


All dissections were made under a binocular microscope in 70% 


‘vor 


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| 


alcohol in Syracuse watch glasses. 
IIT. ACKNOWLEDGMENTS 
This study was pursued under the Supervision of Professor Alex. D 


MacGillivray, to whom I am under the deepest obligations for all that 


| his supervision has meant to me in the way of helpful Suggestions and 
} real inspiration. I must further thank him for permission to use his 
| unpublished morphological nomenclature. I am also ereatly indebted 

j to Professor S. A. Forbes for suggestions and for furnishing a large 


H oumber of Species from the collections of the Illinois State Natural 


History Survey and from the collections of the University of Illinois 


I am further greatly indebted to Professor H. F. Wickham of the Iowa 


State University, who supplied me with a considerable number of spe- 


cies belonging to rare families; to Mr. EB. A. Schwarz and the author- 


fFities of the United States National Museum for representatives of 


seven very rare families from the collections of the. Museum; to Mr. 
). S. Blatchley of Indianapolis for many very rare species; and to 
Dr. Edwin C, Van Dyke of the University of California for a specimen 


of a species of Othnius. Of the many courtesies that Dr. Chas, P, 


JAlexander of the Illinois State Natural History Survey has shown me 


I am duly appreciative. Finally, to Mrs. Elizabeth Stickney, who has | 
helped me greatly in the preparstion of the drawings, I am under deep 
obligations, 
III. MATERIALS 

An effort has been made to make this study as comprehensive as 
possible, including not only a wide series of families, but also a 
representation of the different subgroups within the families. Of the 
eighty-one families , exclusive of the Strepsiptera, listed by LeContef 


pnd Horn, representatives of all but one are listed in this study. 


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4. 
Leng in his recent catalogue lists one hundred and nine families. Of 
these ninety-four families have been studied representing one hundred 
and fifty-five species, of which one hundred and thirty have been 
figured. The fundamental structure of the head is, except in a few 


eases, practically similar for the two sexes, The sex has therefore 


Ze 
been disregarded, except in the case of the brentid, Eupsalis minuta, 


yj the male of which has s long slender snout, as contrasted with the 

| large broad snout of the female. The latter has been figured. 

A number of attempts were made to arrange the figures in a lin- 
l ear series leading from the generalized to the specialized forms. All 
attempts proved unsatisfactory. No matter what structure or condi- 
tion of a structure was used, the structure showed itself to be un- 

| stable within narrow limits and therefore could not be relied upon to 
illustrate a definite line of development. However, the meagre re- 
sults obtained in trying to arrange the drawings in a linear series 
emphasized an important fact: that the various families of Coleopters 
and even the subgroups within the families, have developed along many 
lines. For this study, the arrangement finally decided on, including 
the species,is that.adopted by Leng. This arrangement will be valu- 
able, in so far as the head-capsule is concerned, in showing the need 
for further morphological work towards the improvement of our classi 
fication of the Coleoptera. Owing to the number of drawings present 
ed in this study it was deemed more practical to omit detailed de- 
scriptions. The salient features, only, of the various structures 
are discussed. The following list is arranged according to Leng's 


catalogue, and includes only those species studied:-- 


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COLEOPTERA 


Suborder Adephaga 


Caraboidea. 


1.Cicindelidae. 


Megacephalini.-- Tetracha carolina (Figs. 2, /L4 24§ 37/ ) 


Cicindelini.-- Cicindela formosa (figs.3/2 5/2747 372) 


2,.Carabidae,. 


Carabinae.-- Calosoma calidum (Figs.4 26,750,373) 


Harpalinae.-- Harpalus erraticus (Figs .€74/Z9/64251374) 


Harpalus caliginosus 


3.Amphizoidae.-- Amphizoa lecontei (Figs. ) 


4,Omophronidae.-- Omophron americanum (Figs. 5 272452, 3575 ) 


5.Haliplidae.-- Peltodytes 12-punctatus (Figs.7/238,253376 ) 


6.Dytiscidae. 


Colymbetinae,.-- 


Coptotomus longulus 


Dytiscinae.-- Dytiscus hybrides 


Cybistrinae.-- Cybister fimbriolatus (Figs. 3 /39454 977) 


Gyrinoidea. 


7.Gyrinidae.-- Dineutes assimilis (Figs. 7/35/2535 373) 


Suborder Polyphaga 


Hydrophiloidea, 


8 Hydrophilidae. 


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Hydroscaphinae.-- Hydroscapha nstans (Figs. 


Hydrophilinae. 
Hydrophilini.-- Hydrous triangularis (Figs, S31, a 5G 377) 


Hydrophilus obtusatus (Figs. /4 /32 257330) 
w2 4 


Silphoidea. 


9.Platypsyllidae.-- Platypsyllus castoris (Figs. 


estate 
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10.Leptinidae.-- Leptinus testaceus (Figs./J/33 253 33/ ) 


11.Silphidae.-- Necrophorus carolina (Figs./3 134 2574 384) 
12.Scydmaenidae.-- Connophron fossiger (Figs. /G /3b 260 359) 
13.Orthoperidae.-- Molamba lunata (Figs. /0,/297 2¢/ ) 
} Staphylinoidea. 
| 14.Staphylinidae. 
Steninse.-- Stenus flavicornis (Figs./%/I5 2éZ, 384) 
Paederinae.-- Gastrolobium bicolor (Figs.20 /37 263 359) 
Staphylininae.-- Creophilus villosus (Figs./4 144, 264 336 ) 
Tachyporinae.-- Tachinus fimbrietus (Figs./7/4/ £65 F587) 
Aleocharinae. 
Myrmedoniini.-- Aleodorus bilobata 
Xenodusa cava 
Aleocharini.-- Aleochara lata (Figs. /Z/42, 266, 35 ¥) 
15.Pselaphidae.-- Pilopius lacustris (Figs. ) 
16.Ptilidae.-- Limulodus psradoxus (Figs.2/ /4G, 267957) 
17.Sphaeriidae.-- Sphaerius politus (Pies.22, (44 265, 370) 
18.Scaphidiidae.-- Scaphidium quadriguttatum (Figs.2g VAL ACG 3Y | 
19.Histeridse. 
Hololeptinae.-- Hololepta fossularis 
Histrinae.-- Hister merdarius (Bigs.£44/472/0 372) 
Saprinini.-- Saprinus lugens 
Cantharoidea, 
20.Lycidse.-- Calopteron terminale (Figs.27 /44 AT IFF) 
21.Lampyridae. 
Lucidotini.-- Lucidota corrusca 


Photini.-- Photinus pyralis (Figs.2 U,14- 5 474 354) 


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22.Phengodidae.-- Phengodes plumosa (Figs.2 9/50 7s 39S) 


25.Cantharidae. 
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Chauliognathini.-- Chauliognathus pennsylvanicus (Pigs 30 46/5) 
Cantharini.-- Podabrus rugulosus, Podabrus tomentosus, 
Cantharis bilineatus (Fig;. Z73 ) 
24.Melyridae.-- Collops nigriceps (Figs. /54,27¢6 377 ) 
25.Cleridae.-- Trichodes nuttali (Ties 92/938, 277573) 
26.Corynetidae.-- Necrobia rufipes (Figs.33/54 274,777 | 
Lymexyloidea. 
27.Llymexylidae.-- Hylecoetus lugubris (Figs. 
28.Micromelthidae.-- Micromalthidae debilis (Figs. 
Cupesoidea. 
29.Cupesidae.-- Cupes concolor (Figs. 7% JSS, £14, 400 ) 
Mordelloidea. 
30.Cephaloidae.-- Cephaloon lepturides (Fies.J97 /5¢, 80, Zof ) 
41.0edemeridae. 
Nacerdini.-- Nacerda melanura (Figs 45/9725, #02) 
Aselerini.-- Asclera ruficollis 
32.Mordellidse.-- Tomoxia bidentata (Figs JZ 198 454 403) 
33.Rhipiphoridae.-- Macrosiagon dimidiatum (Sies.@6/99 253404) 
34.Meloidae. 
Meloinae.-- Hpicauta marginate (Figs. 34 (60 284 405) 
Zonitinae.-- Zonitis atripennis 
35.Hurystethidae.-- Eurystethus debilis (Figs FF /6/, ASS FOE ) 
26.O0thniidae.-- Othnius sp. (Figs. ) 
37.Pythidese.-- Pytho americanus (Pics. AS JES 256, 467) 
38.Pyrochroidae.-- Neopyrochroa flabellata (Pies. AQ OZ 257 403) 
39.Pedilidae.-- Macratria murina (Figs. 4 165, 23 &, #09) 


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40.Anthicidae.-- Notoxus anchora (Figs. 4G (ob, 20 Ho) 


Blateroidea. 
41.Rhipiceridae.-- Sandalus niger (Figs. A} A747 FH/ ) 
42. .Blateridae. 
Pyrophorinse.-- Alaus oculatus (iigs. FF A7AU 4/2) 
Elaterinae.--~ Melanotus communis 
Cardiophorinae.-- Cardiophorus gagates 
43,Eucnemidae.-- Tharops ruficornis (Figs. f6//7 274, WS ) 
44,Throscidse.-- Throscus chevrolati (Figs. 45 /6% L7GH/F ) 
45.Buprestidae.. 
Polycestini.-- Acmaeodera pulchella 
Chalcophorini.-- Chalcophora virginiensis (Pics. f7//4A, 2744/5) | 
Buprestini.-- Dicerca divaricata 
Chrysobothrini.-- Chrysobothris femorata 
Agrilini.-- Agrilus ruficollis 
Dryopoidea. 
46.Psephenidae.--Psephenus lecontei (Figs. FE 17h 2ITA | 
47.Dryopidae.-- Helichus striatus (Figs. J7, / 75, 296,417) 
48.Elmidae.-- Stenelmis sinuate (Figs. 54, /76 297 H8) 
49.Heteroceridse.-- Heterocerus undatus (Figs.S3/7/ 275 4/7 ) 
50.Georyssidae.-- Georyssus californicus (Figs. IF (73, A494 20) 
Dascilloidea. 
51.Bucinetidae.-- Hucinetus morio (Figs.5G/77 304 ¢2/) 
52.Cyphonidae.-- Cyphon ruficollis (Figs.54//75, J0l #24) 
Byrrhoidea. 
53.Dermestidse.-- Dermestes lardarius (Figs. 9S /77 30k $23) 
54.Byrrhidae.-- Byrrhus americanus (Figs.9€/3Q 303 429) 
55. Nosodendridae.-- Nosodendron unicolor (Fies. J 7/0 304 44S) 


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56.Rhysodidee.-- Rhysodes americanus (Fies.3 8 Wk, 05 #26 ) 
Cucujoidea. 


57.O0stomidsae.-- Tenebroides sinuatus (Figs.97/8G 30b 27) 


58 .Nitidulidae.-- 
Nitidulinse.-- Phenolie grossa (Figs.66 (04 7422 ) 
Cryptarchinae.-- Glischrochilus quadriguttatus (Figs.6//3530§ge9 

59.Monotomidae.-- Phyconomus marinus (Figs. 63 /6¢, FOF 450 ) 

60.Cucujidae. 

Cucujini.-- Cucujus clevipes (Figs. b6L,157, J16 F3/ ) 

Telephanini.-- Telephanus velox 


61.EKrotylidae. 


Langurinae.-- Languria mozardi (Figs.6é4/05 W4 472) 
Erotylinae. 


Triplacini.-- Tritoma angulatus 


Dacnini.-- Megalodacne fasciats (Fies. CZUGT 3/4, 439) 
62.Derodontidae.-- Derodontus meculetus (Pigs.66/73 W3, 479 ) 
63.Cryptophagidae.-- Anchicera ephippiatea (Fies.67/70 3/4, F#3S) 
64.Byturidee.-- Byturus unicolor (Figs.65/74 3/95 456) 
65.Mycetophagidse.-- Mycetophagus punctatus (Figs.67 Vid, J/6, FU7) 
66.Colydiidae.-- Philothermus glebriculus (Figs./0,/9¢ 3/7 #33) 
67.Lathridiidse,-- Melanophthalma snguleris (Figs.74/75 Ws, 437 ) 
68.Mycetaeidae.-- Phymaphora pulchella (Figs. 7/ 176, S14 #46) 


69.Endomychidae.-- Endomychus biguttatus (Pigs.75/7/ 92g ¥4/) 
70,Phalacridae.-- Phalecrus politus (Figs.7Z/93 324442 


71.Coccinellidae. 
Coccinellini.--Hippodamia convergens (Figs./6/7% JAZ #47) 
Adalia bi-punctata (Fizs.77200 JL 3, 474 ) 


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72,Alleculidsae.-- Pseudocistela brevis (Figs.79 £o/ I24 945) 


73.Tenebrionidae. 
Tenebrioninae.--Alobates pennsylvanica (Figs.7,7 AOL, IRE FH) 
Tenebrio molitor (Figs. BO 203 JL6 447) 
Boros unicolor (Figs. 3( 204527 #435) 
Diaperinse.-- Disperis maculata 
74.Lagriidse.-- Arthromacra aenea (Figs.7/ 205 J25 F#9) 
75.Monommidae.-- Hyporphagus sp. (Figs.%%20G IL7 750) 
76.Melandryidae.-- Penthe obliquata (Figs.37 207 3390 457 ) 
Bostrichoidea. 
77.Ptinidae.-- Ptinus brunneus (Figs.5420% 334 #52) 
78.Anobiidee.-- Sitodrepa panicea (Fies.35S LO? FILA 
79.Bostrichidse.-- Bocstrichus bicornis (Figs. 3€ 24 333, 75g) 
80.Lyctidse.-- Lyctus plenicollis (Figs. #7 Z/0334 #75) 
81. Sphindidae.-- Sphindus americanus (Figs. 37 2/2, 235,456) 
682.Cisidae.-- Plesiocis cribrum (Fies.%3 2/3 G36 457) 
Scarabaeoidea. 
83.Scarabsaeidae. 
Aphodiinae.--. Aphodius fimetarius (Figs.7% 2/4 I37 ATS) 
Melolonthinae.-- Dichelonyx elongata (Figs.Y% 245 333,459) 
Rutelinse.-- Pelidnota punctata (Figs. %Z 2/6 FFG ZEO ) 
Dynastinae.-- Strategus julianus (Figs. 73 217 G#O #G/ ) 
Cetoniinse.-- Osmoderma eremicola (Figs. 9732/9 oH, £62 ) 
64. Trogidae.-- Trox suberosus (Figs. 76 2/7 JFL, 47) 


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WG WS, a8 aed jixxagld whee ie —ni 
ete s\ 5. Ve ride bi fourgneks ait oqd amg al 
Cit» CEE +S. * Grass “aot nT At bo yuh iS .<- «e380 Zt 


‘ Aye a 4 o eS ; Si *. 2) al [ ff CT") 7 4 toqsgali =~ 
Y he w 


hire TES, . We icedt ds) spent 1u Liisi ees 
SNE SR ont) aeeyeeie ayaoledoes ~« BAAS oF z 

: r WS SR.2al eh), eronvy aioe LDCR ~~, 09h t@ 
» 6 o828) ernetlot sopetatee “+ saenttant 
e& @\. le nal€y ale 6 bwe ts enteboni a», Aen 


pt RX ENS. WE eget) svaoiedus. zene me of 
(yah tat poo ‘Xi ogi!) emek echephepeet be 


86.Passalidae.-- Passalus cornute (Pigs. 221 344HS $66) 
Cerambycoides. 
87.Cerambycidae. 
Prioninae. 
Parandrini.-- Psrandra brunnea (Figs. 0S A224, 34S 467) 
Prionini.-- Derobrachus brunneus (Pigs. 223 3 #6, 265 ) 
Cerembycinae.-- Glycobius speciosus (Figs./&¢ 224 747 £59 ) 
,Lamiineae.--.Tetraopes tetreophthalmus (Figs.MZ42425 345 £70) 
88.Chrysomelidae. — 
Donaciinae.-- Donacis piscatrix (Pigs. 642246344 Z£7/) 
Orsodacninsee.-- Syneta ferruginea (Figs.77 227 JSG, 47-2) 
Criocerinae.-- Criocerus asparagi (Figs.W4225 351 47—7 ) 
Cryptocephalinae.-- Cryptacephalus quadruplex (Figs Mz1yIZ74/A 


Eumolpinse.-- Chrysochus auratus (Figs M/ZH3535, 476 ) 


Chrysomelinse.-- Leptinotarsa decimlineata (Figs MELY TI V7) | 


Galerucinae.-- Diabrotica 12-punctata (Figs M25 355 47F ) 
Helticinee.-- Blephsrida rhois (Figs. Q Z38 356, £75) 
Hispineae.-- Anoplitis nervosa (Figs.// AVF W747 7) 
Cassidinae.-- Chelymorpha argus (Pies MZ 235 353 957 | 
89 .Mylabridae.-- Pachymerus arthriticus (Figs 3, 23607 7 4-80 ) 
Mylabris obtectus 
Brentoides. 
90.Brentidae.--Eupsalis minuta (Figs M7237 Jee $SZ, ) 
Curculionoides. 
91.Belidae.-- Ithycerus noveboracensis 
92.Platystomiidae.-- Eurymycter fasciatus (Pies MS LIF 36/ $F? ) 
93.Curculionidsae. 


Rhynchitinae.-- Rhynchites bicolor (Pics MZ L3G 36ZL4%4 


ae ARE *A, 


i tae a® & BAS. 2 


Susy OeY WSS Lae FE) 


\ 2 1 oie é 7" — Ws = 
ah UG FS ) 
A SA AOU Bas 8. ced 
Bays. y= eA, ~~ > + 5 
J > 4 
Y z > . x. Wa 
.% L< * . at ‘ ¥ 
‘af > wis : 
‘ 
LV Sot , d 
[ » d am a 
he So) HS i 
é ix i 
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if 
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Kee Gay ati. aXe 
1 i * 
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{(- 33) Den 


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rh vt 
‘ie he “OS 
Ai en | 


yee 
‘ime © 


volt iedita, sewenipieas Se 


ye 7 
de eo 


7 


ecnwrTe rasta 4 


are ae soactoam yal 


Meta satay 
igek 5 eg toate ot 5! 
{sr t ao lt ainda ~— 
Tt) atons sidgedy Ey: 


4217) atovren alee S 


| 5 an 
4) copce’ uesompiomy aes 


i 


ses a bimaigie 


oe?) et doses 


} 
i — 


a , 
: : 4 


J sacossy dakam SaRpaoganh o6 
seul rot ogageee ~~ +z. mo 
hie 


| 
I 
' 
t 


MH 


12 
Attelabinae.-- Attelabus snalis (Fies.0270 HZ43S5) 
Allocorhyninae.-- Allocorhynus slossoni (Figs. | 
Otiorhynchinee.-- Epicaerus nubricatus (Figs Me L4/, W674 FIG ) 
Curculioninae.-- Lixus macer (Pigs./23 242 56€ $55) 
Thecesterninae.-- Thecesternus humeralis (Pies M7 LFF 367 457) 


Calendrinse.-- Sphenophorus sequalis (Figs.2/ 244 56 5497 ) 


i} Scolytoidea. 


94.Scolytidae. 
Scolytinae.-- Scolytus quadrispinosus (Figs SLA 2 4S ICSF HO) 
Hylesininae.-- Dendroctonus valens (Figs.W3 2g, 36 7A7H) 
IV. HEAD-CAPSULE. 

There exists a distinct homogeneity in the general character of 
the structure of the head-capsule of Coleopters. Its uniform strong ! 
chitinization is typical. So is the spacious area occupied by the 
mouth-parts, producing a relatively broad cephalic end. Especially 
characteristic is the wide space between the occipital foramen and 
the submentum. Then, there is that indescribable similarity of struch 
ture, even between groups widely separated, that can best be appre- 
ciated from a thorough knowledge of the morphology. As an illustra- 
tion, there is little superficial resemblance between Harpalus (Figs. 
625/5%) and Phalacrus (Figs 7Zsywée, either in external or. internal 
morphology. Their distinct differences are merely due to two diver- 
gent lines of development. The structures of the dorsal surface of 
Phealacrus have become highly specialized, whereas those of Harpalus 
are relatively generalized. On the other hand, on the ventral sur- 
face the metatentorina has remained in a relatively primitive condi- 


tion in Phalacrus, whereas in Harpalus its position is highly: specialf 


ized. The internal structures of Phalacrus are rudimentary or lack- 


ics: ant Cae ae ethnns oes 
‘a _ -G2 ft) “Eisvecte »regtiedaeee ba 
Di. Get) artgoltedb: ay su tee nar 
(et DU cS. EEN, 11 » er FE SAE 


4% ett es.ek, iol ud a mr cit } . oe o @ Tae ya ae 


x. 
CERMAK ADEE) ate ‘deb godeuenee 
3 : ane ry >» \ dave ; a z 4 : ‘ = 7 ’ “> 
- © * 
i ies , Ta 2 ) — ‘ah 
cya M4 
' % > a AS Dui 
‘ 
¢ 2 
. 5 eva 
6 rt cone saya 


anc) TY sl a =a 1 «SS { Ao = _ eats aay 
| fe yee a 
i “45 t =u hy f ‘a <f2 saggy’ Fa 


i é aiqs 2 salow 9 biy: 
fa j é -on\ie Sierra tem edi 
= ) ess 


qos (fehid Sasori aiawee 
eerteoliy me ) pnt aderoee tant dues 4 
eit) nxt feo soba fave, beeaie oat anient = 1 
bbe fe i vec? Be ~, Wee sv eat niSaqnee a 
‘ | ; m2 geons teeny Ea 3 ted * 
v2 ‘ he ent orenyetewob vo* 
elisa Seer be boa? Lottaes iia eyouss att 
jlian.? Deneay agit eo omstatenes wie 
=i evi ) fe vi¢elcr es of Denbanei eae actfeationd oO 
: wl hesy ot. Saber ae lease 
| aS¢ai 1° \talnesniset ean Satsaiads Be sernloetty 
ee oe 


| 
| 


/ment of a compactness of form of the sclerites that do not consoli- 


13 


ing, while in Harpalus they are in 2 well-developed primitive state. 
Considering the degree of generalization of each species, there can 
hardly be any question that Harpalus is the more generalized. The 
above comparative description simply illustrates roughly the problems 
of complexity of development that are encountered. The two distinct- 
ive kinds of development as shown above for Harpalus and Phalacrus, 
that of specialization of the dorsal surface and that of separate 
specialization of the ventral surface, do not in the least necessari- 
ly parellel one another in the Coleoptera. Indeed, these two lines 
of development are predominantly divergent. In the majority of spe- 
cies, the development is trending towards the obliteration of sutures 


and consequent consolidation of sclerites, and towards the develop- 


date. The general trend towards a cephalization of migratory struc- 
tures is a part of this process, too, as well as the development of 
&@ stronger chitinization of the head-capsule as a whole. The entire 
Phenomenon appears to be fcr purposes of strengthening the head. Be- 
sides Phalacrus typical examples are Tenebroides (Fig.45¥), Glischro- 
chilus (Fig.é/), Megalodacne (Fig.65], and all the Scarebseoidea. 

On the other hand these processes have lagged behind on the ventral 
surface_in Phalacrus and others. It is true that the hypothetical 
type (Fig.Z47) shows a marked cephalization of the submentum, with 
obliteration of sutures resulting from this migration. Yet the pre- 
dominant condition of the heads studied shows only a certain degree 
of removal from the primitive type. Stronger chitinization has 
probably kept pace with the cephalization of: the dorsal surface, as 
well as the growing compactness of such a structure as the submentum 


But the very significant structure of the ventral surface, the meta- 


ta, ov it inficG’ Seno terabe Tien: 2 al oxE meat! ; 
, : red? ,saeinege nde Te 21 tps Ciao 
; 262 .ise¢feretes eron er ,2: ss Saree 
Be dera ot? gis ‘Ll 2ecs wren a te sree 
Besuitel> cut eft .hatetacosan. ote raat? PROS: 
oe pOartt fl toe 27iayi7et rod rt foendge 
| etateae: , | | pe 7.08 oct te gots ; 
i 2 Oe, 7: oo : Miah Lasihawe say 
| os ie) off ak caditons, 


“vou 


t 
, 
= 
ty 
r 
Lan 
Lie 
a 
Lg 
‘ 
> 
S 


jjeo = afqaned hnomm 
Behe. tee hd | a’ of ,00? eeapkg hae 
} 

ee i 5 - DASE ag ~0 ‘aout 
aoe jirh a: ae ine areetys ‘Te cece Tee 2 ag et, 


prises: ARSE OH) cabreedel oF ace cane tne 
; cum , Rew ecanbetepen 0% , 

ez> mer ad fed Songet "vee esate mead } 
bar Faonca 4 edd pax nd zi orate ” ] 
eeinendsu ad’, te oa itaytlodeee eatin a atte: 

ve potters io. 2207 nods wo cune aeons, 20% 

atte ' cinu oworte besbedg sheet axit 30, <4 cele 

H oolvarttalel.g +seenerts (equd ouiiiiaias ose nie 8 

prises Coezeh fy Se opkhenhindght: am? atte mean Bei 

, eceerr | ee siatphtie ¢ Gon te daend GpgK0o, : 
“ied <7 ,cottau Lewneyr ait Se etebpaeth?. 


14 


ina, only shows a certain degree of removal from the primitive con- 


dition in most of the heads. 

Developmental processes such as are discussed above can be best 
worked out through a comparative study of a large series of forms. 
In fact the determination of the homologies of some structures enter-| 
ing into these developmental processes, as for example the various 
changes in the epicranial suture, and the determination of the nature] 
and line of development of the ares between the occipital foramen and 
the submentum, would probably be most difficult without this compare- 
tive study. The chief value of this study lies in all probability 
in the determination of homologies, to the end of understanding the 
lines of development present. The homology existing between the var-} 
ious structures of the head-capsule of Coleoptera and other orders of] 
|insects, particularly the generalized orders, seems to work out satis 
factorily. From such an homology the hypothetical type was construct 
ed without much difficulty. In general appearance it is oblong and 
rather flattened dorso-ventrally. Such a form is fairly character- 
istic of generalized insects and of the more generalized Coleoptera. 
The mouth may be considered as directed cephalad. Such a direction 
is representative of the vast majority of the heads, and for purposes 
of convenience, at least, the following discussion considers the head 
as extending cephalad. ‘Some possible exceptions in which the head 
appears to be directed ventrad are found in Calopteron (Fig. /#9), Ma-| 
erosiagon (Fig.f7), Tharops (Fig./7%4), Throseus (Fig./6¥%), Byrrhus (Fig 
/7¥O), and a few others. It should not be forgotten, however, that in 
primitive insects the mouth is directed ventrad, and the occipital 


foremen is on one side instead of at the opposite end. 


Epicranial suture.-- The line of closure of the head in the em- 


% 
S #8! 

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enn oa uApA a 8 22 OeRs - BOGOF * —_ 
gee opaurl e ‘Bute err sor) ® = 
wg.t moe fo selbyeolonen iy, dont 
aljears 4 Osasuna Ket 
rls awl oatthe ted 
arly te der? ai . ad ‘Be: 
Ya teh Peta oe videdang t 
ti) to outer? 
7. ig ie Combat he 
CLince Cae | Seeman £ 
ee ee ee aoa > aba ‘ett 
¢ “eed te ~onmy one 7 
5 ait qrotammty, he veoh 
a sco 
— Hone Jee cevtewnoanah” 
; eters O40 t to bee, OPpeeg eh 
Sateudue Drees: oe 
. A ae Dem tpt dade 
; sof) syhiow 7am ais mote 
ote ack ‘geome elclouee oot 2be 
}. a vt iy. igv tJ aie Say hooey, > 
ev eee) ace “ae (Mi, at) eqprad® , ee 
ay pettosigt ad rap) higowdks 2 cowie 
ns y haat aoe ied deel ol aeons ve 
tue wtteog@e oth ra Xo Tinodint 2 
462 60? Te gle % apmkt eit? ‘fh £ 
= J2. = orn = es is - cE 


15 


bryo is represented by the epicranial suture. The complete epicrania 


suture is typical of generalized insects. Where it is present in Co- 


leoptera, this denotes a generalized condition. The primitive form 


of the epicranial suture is that of a deep inverted Y, with the ce- 


phalic ends of the arms near the lateral border of the labrum. The 

ee head is represented as having a complete epicranial su- 
Iture. The epicranial stem extends to a transverse line drawn through 
- middle of the compound eyes. Branching here, the epicranial arms 
continue to the margin of the head cephalad of the compound eyes. A 

——- epicranial suture is not of general occurence in the Coleop- 
tera. It is practically complete in Hydrous (Fig.f) and Hydrophilus 
(Fig./2), 


So in the former, and characteristic in form. 


very distinct and sharply invaginated in both, particularly | 


Each arm reaches the 


margin of the head almost immediately cephalad of a compound eye, and | 


(Fig.d 34) 


the arms are not as generalized in position as they are in Epicauta,, 


where they are quite distinct. The only other occurrence of a com- 


plete epicranial suture is in Chelymorpha (Fig.y2). Here the arms 


meet the stem farther caudad than in the other genera named. The arms 


in this last genus are distinctly curved, as contrasted with the more 


or less straight arms in the above-mentioned genera, 
The epicranial arms or some portion of them ere present in all 


Coleoptera, except possibly in Calopteron (Fig.Z7)and Photinus (Fig.3) 


One or more species of every superfamily of the Adephaga and Poly- 


phaga, except the Elateroidea, Byrrhoidea,Rhysodoidea, and Rhyncho- 


phore, have the arms complete. In the Caraboidea they are prominent 


= 
os 


S$ nearly straight sutures across the head, as in Tetracha (Fig.Z), 


icindela (Fig.3), Calosoma (Fig.¢), and Harpalus (Fig.@). Their 


most generalized condition in the Adephaga is found in Omophron (Fig J 


icp fants ices 


=: 
| 
4 
: o 
| 
| 
5 
| 
» | 
hes 
a 
' ae 
d, 


oe 


at 


An” 


+ 


t 


ef 


Ye niadp im ‘st qelew #3 ~fdineoy oe 


spt ‘at cpactOhs basal ai sot 


ger =4  ~oavnere Uitenteg eb ath 


ie a av 


iy 
& 


,f6 Beker 


a 
i 
. 
. 
’ 
sf 
4) 
=" 


af aigen Wiad sihebute 


ts tees nh: Bee “nme 


lair ohh Sanhite th 


>) : 


Ke tomgt ote | ao ae eset ag 

Tar y. ee is BS os 4? babar ctv 

or Logolt edge att Af émsa dh 

; be ne Pat) ca oe 

ds tv arittog anee 23 aaeee 4 

aLigal fonwe Yieve to 46 i 

ith , ah Leabieaes sob lovetols. ost 

bistered oak ge st otymoc gate aut 4 
beod o¢t s70nee Renanae re 

7 vin 


oa ine os 
tape 


AP on 8g 
¥ > 


16 
in which they extend from the meson at a sharp angle. . Representative 
Species of other superfamilies that have the arms complete sre: Ne- 
erophorus (Fig./3), Tachinus (Fig./7), Chauliognathus (Fie.3o), Cupes 
(Fig.J6), Cephaloon (Fig.37), Notoxus (Fig.#Z), Heterocerus (Fie.52), 
Eucinetus (Fig.50), Mycetophagus (Fig.é9), Tenebrio (Fig.3O), Bostri- 
chus (Fig.jG), Aphodius. (Fig.90), and nearly all the Cerambycoidea. 
Species having parts of the epicranial stem preserved are not very 

icommon. In Omophrom (Fig.5), Tachinus (Fig./7), Penthe (Fig.J53), 
a & number of the Cerambycoidea, parts of the cephalic end can be 
jidentified; in Omophron (Fig. 5), Phengodes (Fig.297), Cupes (Fic.g), 
ISitodrepa (Fig. JJ), Blephsarida (Fig.//), and a number of the Rhyn- 


Bie. parts of the caudal end are present. Chalcophora (Fiz.¥7) 


jretraopes (Fig./% sre peculiar in possessing practically all of the 


The arms in Chalcophora are as short as 
in any other species studied. Parts of the arms are present in every 
Peer of length from nearly meeting on the meson, as in Nosodendron 
\(Pig.57), to almost complete disappearance as in Chalcophora (Fis .Z9g) 
jend Rhysodes.(Fig./7Z). They also show varying degrees of disappear- 
ence and invagination, from the deep, distinct invaginations of such 
forms as Dineutes (Fig. Y¥), Necrophorus (Fic./3), Tachinus (Fig./7), 
Heterocerus (Fig.JJ), and Arthromacra (Fig.7/), to the faint or slen- 
her and shallow or not at all invaginsted sutures characteristic of 
he Scarabseoidea. The character of the invagination associated with 
the epicranial arms is not as Simple as may be thought. In Harpalus 
Bigs.674 44), the epicranial arms extend from the meson along the 
pdge of the invagination to the pretentorinsae,from which they extend 
o the bottom of the invagination, curve laterad, and continue to the 


hargin of the head. The course of the epicranial arms can be better 


Pi thaiacsevge. wstsox yoke es Ye -nceedl Ger meen - 
(eons HAS ct) Sent Teds ww Lema i 
ae 
Dieg om et, 8L87 odplived) ..( Nie et) npn) 
Deeg te) acdsecoxel! , Giy.a2k) oxet ol en 
ioar (Wet ) ofiiome t ~! te ae 
beds Of a ev Te! 2 aie. , 1. oe) “ c 
| n° ta bev fo4s= : igy 1 a 
i lek. ) i <1 a % rar —tZ) 1 
5 On | ome omnacnet “a 
AS ee | VS 12 ae co 
| nist 2 {xs {tinct ted 
hue fi) evangootsd s 2 iaepag att 7 
/o 
bai! te 4 i tft og 
{ ar ' ; i S| ~* alee aah % 
i Sr é ; une detours 26, ad | 
Oo efies ale pee “tsbon meat ¥ 
A 
| tue*.. < TH6 < ie stwlquoe ‘ 
- _ 22 ~9 6. sv rods sé i eat e ' a 
: | arfsnib 2ags acd aot not? 
its B\.0 27) sunodtadehbelt. | a eee 
sat of Vere af bas - (phew 
2 horn t i ft aq f val Life 28 ton 64 
aly -y 4o¢ebands egT 5 
(mene id ei 2 asten st ieee 
72 Lotaarerge 6a, 


17 
understood from Omophron (Fig.S), a related genus, which hss re- 
tained the cephalic portion of the epicranial stem. The line of the 
invagination appears to be and often is considered to be simply the 
Clypeal suture. If a specimen of Harpalus is soaked for a long time 
in potassium hydroxide the invagination can be opened and the various} 
structures in this region studied advantageously. The invagination 


when opened (Fic./67) will be seen to assume a deep wide wedge-shaped | 


form, extending entirely across the dorsal surface of the head. The 


pretentorinae are located on the external dorsal surface just caudad 
of the cephalic margin of the invagination, Although the epicranial j 
arms are not in evidence anywhere between the meson and the preten- 
torinae, it is assumed that the cephalic marginal ridge must repre- 
1 sent them, since the pretentorinee are not only located caudad of 
the invagination, but the arms are quite distinct, extending from 
the pretentorinae to the bottom of the invagination, in which they 
then curve laterad and continue to the margin of the head. From a 
cross-section (Fig.7f) it will be seen that the pretentorium expands 
cephalad in characteristic form from the pretentorina along the epi- 
cranial arm: to the bottom of the invaginetion. In Calosoma (Fig.¢ ) 
the seme condition of this region is found as in Hsrpalus. In Omo- 
phron (Fig.4) the epicranial arms sre distinct between the preten- 
torinae, extending from the meson along the cephalic border of the 
invagination. From figure/6¥it will be observed that the cephalic 
porder of the invagination in Harpalus is along the imaginary line 
of the fronto-clypeal suture. This border may represent the cephal- 
ie limit of the front. The invagination, then, in Harpalus and Ca- 
losoma includes the entire front. The line of the invagination in- 


steed of being solely a part of the epicranial suture is in fact 


i? , 
Tt 
Ls 
ri fy oO 
ie] 
eet 
_ 
' mo 
Aa 
Ss We 
‘ 
‘ r i 
| 5 
{ n 
a aw i 2a) 
7 
4 
f >. 
, 7 
) 


lawia of oF Ker ghienem Bi nahie Hae od be 4 
ae y & , BolT oc I mii yo i TD ‘4 oglne oa a a 


emisove f ay eT Bnet es oarhe i Poqts oes 


—, ert te rey. 2 ee) ae oT va 1s seamed 


i) 4) a> a he 1. . A a] al | ‘Fe a tr ae ete z 


uo dio  ponek Setaten- 2 47 ater seneen 
pit eit. sete, Paloatotes ons te nothtod, 

a ct & a lef lv A ej batt : Og (a0 ie “wl ont | 2) 4 
VaR ah : iT, A ‘ Ou jehs et Lisa 


“0t tnardiice Sibae baracot “ana 


nT 2! om ’ aft Phe lies 


bars witeih tine lage aay ‘pat ce 
i L obra Raval ars Fe an thea ot or 
or siaetied aoa’ Det 


Sn wf say yey ee, CDee te Rey 


wo) 1 aul joe Deer oltetsed 
ad | LOL Saher SR To apitod elit 

lov.) ab otet SRR te oot 

bie we.4 :tntaeiigs pate 
rod pifaives sit decte moe vn 2c) aot gr k 

1103 aid forieado oc {ile AP eee aoe 
a) war eanteed nclagrellgt shoal avae: 

jot Get. cabiod @4) » eae 


on! hit  hebhihyege eee sage itt 


9 


= 
mt 
; 
| 


lve ek 7.8 1 to ee2a ea? weort aut 


al satuson let paepege sat SO 2taq 


18 
compound in nature, representing the approximation of the caudal bor- 
ders of the front and post-clypeus, and that part of it between the 
pretentorinse may be termed the clypofrons. . lLaterad of the pre- 
tentorinae to the margin of the head the line of the invagination is 
readily seen to be an approximation of a part of the vertex with the 
caudal border of the postclypeus and cannot be included in the eclypo 
frons. Due to the more primitive position of the epicranial srms in 
| Omophron the invagination in this genus contains only s part of the 
l front, hence the line of the invagination between the pretentorinae 
} is Simple in nature. In Tachinus (Fig./7) the epicranial stem ex- 

j tends distinctly into the invagination, the arms continuing in the 
same to the margin of the head. As should be expected, the preten- 
‘torinae are within the invagination. The line of the invagination 

# in Tachinus is then of a different character from that of either Har- 
palus or Omophron. In Tachinus, it hes nothing whatever to do with : 
the epicranial suture nor with any other suture, being throughout 

the approximation of parts of the external dorsal surface of the ver- 
tex and the front. It is obvious from the sbove discussion of three 
types of the invagination associated with the epicranial arms that 
the dorsal surface of the head-capsule in Coleoptera must be studied 
most carefully before a correct interpretation of the parts can be 
made. This is most true in the case of any invagination that may be 


present. The latter may not be readily observed when the head-wall 


is strongly and darkly chitinized, necessitating treatment of such 


specimens before the parts can be clearly made out. In some genera 
such as Dermestes (Fig.§55), Scolytus (Fig./@Z, and Dendroctonus (Fig. 
Ag) 911 external trace of the line of the invagination may be lost. 


In such cases a true understanding of the parts can only be gained 


Bil aati dah oS is pa ree i RNs 
. if | t | 
i-tod Lolusn e4¢ te sodtagbesugqe edd anitReneges 
ais Hoowed 1T£ TO Oeq) Ree OAe arent lovdnait 
| tq ate To boerereg” 7 jam TR rete sit Samed -e¥ 
; iolseve: at Ye Gest ede bagd as? ‘to aired 4 vd 
ee Hn 

od fie xertav cee Te Pang = Le ot Heamtxesgge tia — 

} 
Boy. ie at Delieioad we sonmeo hae auagyivsaeg) eat 

Peni secs fafaroiqe ait To eottleoq Svid imi'tg snipe sa 
| Bi r 2D PSag) a Pane tasefoge Giteds pict aia ogleent | 
Ae ixotnsretg at? aeedted noLtedinevad out ts “oath visi 

«246 mote fe bie qo of% tN sid) eaninna? a at 5 

5 ui ‘*l timivnoo mrs edf. ,Aoivanigavar-egt osu’ 

} nébe. ‘ af? Oo? ongte od Pivote oa been ot ork 

iri. teats ‘ed? +o. eBlt ory’ .tolvs ohgevat edt — 

® wm \edtie “db tose west saad et a doete rssh 4s to. coat 

| atic ab ct reTeva7 aches ry a4 sentdoet al 5 ie 
; Pirodsdort? naiga . “iatyve weita qe aithy toh ROVER 
j "a ips 


Sigedndy *%. saolezzoeth avoda adi moet seutvde ef 42 r) 


flaw-lcat adfY sade Depress do Glabeee ed-Ton Gam t3errel 


tenes wand ol rvo obam vlveete’ ed os6 Straw att erotel 


ac? amis fetnetotce eft ddinw Bete facaese apiT 


6d Aso ® ; ‘ea a ao hited ongaetisi foertog ee tghed: 
fose lo taentees? goltatigeooen \Seslatt ity yitees Ban £ 


a .da0 ¢ var re isan tyates ent’: to ambi: oth eaters r 


Te” te | fLeniah Lantedee oad te ariay) TQ) 


ctet.osfe® wt ‘wlaiasdeboced’ 4ff to 


sf 
S 
i 


: ag ea? coltteucinveval yea Dol eaas 4? cf oot? Gao, 


\) weno sorbte€ Dae Ret) eatphdosg( Th.geey 


ties b4 oleo neo whaeg ond + eeneae 


, y is u : we | ri 


18 
from an examination of the ental surface of the head. But in aes 
ized forms the ental indication of the invagination may also be ef- | 
faced. 

The epicranial suture can always be iocated from the determina- 
tion of the position of the pretentorina. The letter is always close 
ly associated with the epicranial suture, being present either in or 
just off of the suture, in which case the pretentorina resembles a 
sort of pocket. There is usually little difficulty experienced in 
locating the suture. The cephalic ends of the arms sre the most per- 
sistent parts of it, being present when the remainder of the suture 


eannot be identified. Interesting examples are found in most Rhyn- 


chophora, where the remnants of the epicranial arms are represented 


by short furrows located at the cephalic gnd of the snout. The epi- 


|eranial arms are typically structures of the dorsal aspect, but with 


the shifting and modification of other parts of the head may be con- 
fined to the lateral aspect as in Dineutes (FPic.fyy), Helichus (Fis.d7) 
Adalia (Fig.z2#%), or to the ventrel aspect as in Cybister (Fig.z25), 
Hydrous (Fig.7%), and Phalacrus (Fig.3z). 

Vertex.-- That part of the hesad-capsule not embraced by the 
three primary sclerites cephalad of the epicranial arms, the occiput, 
and the postgena, constitutes the vertex. Its extent is determined 
by the form and size of the three above-mentioned sreas, For in- 
stance, in those species with much reduced epicranial arms, as in 
Creophilus (Fig./¢g), Limulodes (Fig.Z/), Adalia (Fig./7/), and Phalac- 
rus (Fig.7/), the extent of the vertex is correspondingly increased. 
In the Rhynchophora, as represented by such species as Lixus (Fig.s3) 


and Sphenophorus (Fig.&#/), it is very extensive, including practicel- 


ly all lof the snout of the dorsal and lateral surfaces). The area 


+ Beh ee = 


- tate GoreS ptt ae ah re 


Piet ove nf St | yhoott ath, hO ecatrus inne Sr 
S =7* ed ovie Yao cetéon byt! Sit ko sole tias¢ 


shitsitaetoh aif moet Delete: of aeyouta ran’ Be. iad 
meotiac agoeria wl iy eit te Fa Ly oe j 

ca ok: Heed te Seeeene Wile’ .aruit lehetene 
i a seidmaues eats | O68 ye plat ‘ Lm et KOREN S 
thi afit wii is we are 


+} 


wor fuox ott ote Shes iia baa ites ig8O out . SR 
} - my) 


5 

if 

' 

: 

y ; : 

y tus? g 
oe» 4 i oy a ’ ee 4 ej be 
St7 wih 3 Ce { 2 \ a 7 
tia 

4 J J ; Ay 

oe “evi. toce al Cami Sizer cb! se eee 

i = ; A 


hodan on rug >s woh M ithe “ <p I a4 I a - i ee #Y etry Ls 


Deity oc Bost Loh! i yar A td 
Pex. ahs) eatet iat of. ik ttl Sy~ 6a $ conga ¥ 
i « 7 7) gege hb: tosbgefy LETTEEY weet on. 
H AST. aa) nancial Dalene. is 
Beactoon of , ewes eee! i oo §otesgeoteiiiess 
bentogeteh of knots 4). > Seeder one nota eae 
rs <fee18 fer nebeevose: seudk’ 2ae to, enna: 
hoo . dene Veblneavldé send iee ulboe Dim haan 
om! bh = “s “ee ee )') tan y = PT) 060 kay 
, hous ef «denthoeodharten al eerdey ath as 4 ott 
alt) Kar » 2elvege vohe YP Ae eesetgan 
‘. pl? owiq:ani wot. , ovlsagiice: Fryer es itis 


ehus ot? ‘jovavhii Ueieg eae a 


located on the latersl aspect cephalad of the compound eyes, between 
the latter end the epicranial suture, is the gena, a part of the ver- 
tex. The limits of the gena are not definite. The prominent ridge 
in many genera, dorso-mesad of each gens and antacoria, is the so- 
called frontal ridge, that extends in the general direction from the 
epicranial arms to the mesal margin of the compound eyes. The fronta 
jridge is prominent in Harpalus (Fig.6), Necrophorus (Fig./3), Tri- 
Hchodes (Fie.32Z), Pyrochroa (Fig.#0), Dermestes (Fig.3$), and many 
others. | 
Occipital Suture.-- In generalized insects the occipital suture 
#isS confined to the ventral or caudal aspect, beginning near the lat- 
eral margin of the postcoila and extending around the caudal or dor- 
|Sal margin of the occipital foramen. In the Coleoptera this suture 
Harises laterad of the postcoila, extends cephalad for a considerable 
| distance, then curves abruptly laterad, extending onto the dorsal as-| 


pect of the head, where it joins the suture of the other side at the 


| meson. The genus Cicindela possesses the most generalized condition 


of this suture found in the Coleoptera. The cephalic end of the su- 
ture is modified into a ridge. This ridge is considered a later de- 
velopment, and is not shown in the hypothetical type. It unquestion- 
ably represents a part of the occipital suture, and can be identified 
in practically all the Coleoptera, as in such widely separated groups 
as Molamba (Fig.26/), Nacerda. (Fig.Zg/), and Byturus.(Fie.3/3}. The oc- 
cipital suture separates the vertex from the occiput and the postgena 
Only the Caraboidea seem to possess with certainty an unmodified oc- 
cipital suture. In Cicindela (Fig.3 ) it is complete; sand nearly so 
in Tetracha (Fig.4), but very faint in great part. In Calosoma (Fig. 


250 the unmodified suture begins farther caudad and is more distinct. 


ye) 


i] Se a, : a 
H int ttt we Cecio ev TOKE LATO Foojes? ata i 


Cotmie ghye® 


6v ais) > ee ria Ve 
- 3 Geek i ; ater, - oat 2BG, 
) ’ ane » SY eno to fe&60ne0R ye 
; iti : Laers + Ae ¢ 
7 £ ie “+ f » ss i 3 sig ast 
+i #: ’ « 4% ‘ . 3 . ; t} [agian ne. 
{ Ne ; 
= 
| Ce ) - m4 ’ > . a % } 20th oot 4 a 
aa in 
e > 4 > j f i Be ix ‘4 ~~, Ff 
13 f r ¢ , 4 r af naw 
f 
( \, | 
© toi Faber if ho tas Cf ORberae 
i ~ ; a a ; 
i rf : Ube Ayo ‘ cs M4 se", E E ‘ Pes i . 
i : Le bs y 
4 ¥ fii a ii § BOC . «i > ’ 
- ? a ¢ 2 54 
6 -ehte if «¢ Digit) Ge 
' i. Ce 
a os i> eirbor, ve cet 
m tao ccateeot afgh te 
! * ; 4 ’ tf ‘ot 
i . 
z oul re a bam 


core: CGRP 
tote 


fa 7 ¢ 
yw wou ae guadyoafed exit. 
, REF ort ine Nee) 2h 
swwloot. get nowtk Zee tee aa? vetar 
: ry st 2k ek SES » 1 bade O-A ie 
eoseied ni .fted Feet kat €x@e tit r 
- ; toa [ heA 2 4 = ute et mate ‘ 2 
ae s.. e —— — = 


3,4, 


21 
In Omophron (Fig./27). two short, cheracteristically curved, lateral 
ridges no doubt represent remnants of the occipital suture. The ridge 
across the lateral aspect in Peltodytes (Fig./29) msy also represent 
ithis suture. In Cybister (Fig.259 it is probsbly represented by the 
crescent-shaped suture on the ventral aspect. Ridges or furrows ap- 


pearing in the same general location in other species, such as Aleo- 


Wchara (Fig.2é6), Throscus (Fig.293), Cyphon (Fig.30/), and Aphodius (Fig 


1337). may possibly be homologized as occipital sutures. In most cases 
these'ridges seem to be merely to mark the limits to which the head 
is telescoped in.the prothorax. 

Occiput.-- In those species possessing an occipital suture the 
occiput is recognized as a distinct area. It includes the region be-| 

I} tween the occipital foramen and the occipital suture as far as the 
postgens, appearing as a sort of broad band across the dorsal aspect, 
divided by the epicranial stem and indistinguishably fused on the lat 
eral aspect with the postgenae. ixamples of a well-marked occiput 
are present in Tetracha (Fig.4), Cicindela (Fig. 3), Calosoma (Fig. 
4), and Harpalus (Fig. é@). In Omophron (Fig.Z7) the occipital su- 
ture is so short that the limits of the occiput cannot be definitely 
determined. In those species not possessing a recognizable unmodi- 
fied portion of the occipital suture, the limits of the occiput can 
only be judged accordingly. Even in generalized insects the occiput 
is nearly always fused with the postgenae, and is so represented in 
the hypothetical type. 

Compound Eyes.---There is a great similerity in the form and lo- 
cation of the compound eyes. The general form is oval. They are lo- 
cated near the middle of the lateral margin of the head. Such a form 


and location is given in the hypothetical type. There are a number 


[oye Lag 


me 


— 


oT) 
ia 
4 


beveto vJ Taniéeiretesteno. . sede emt. ie 


SER PSG SAT To ahiaeeg abve sea 


+ 


‘} .carykovle 


i RS. wit) saree 
Ie .t nev ef, 4a otk 
BOL Sol 123ao 

5 
LF X out ae ni i 
= 
ee ae | aly LSiTtes?g 
. I t 
{ Sere 
c if C Sie 
: i C2 2s 
2 = = 
a . fatness 


serene caceate 6g are 


“5 ; 


=r: 
ay i 
oa 
[ 


—+ Ry) Eo 2 es 


Ad 


of interesting variations in form from the normal type. Dineutes 
(Fig./33) and Tetraopes (Fig.soz) heve four complete eyes. This phe- 


nomenon is produced by a projection of part of the vertex.into the 


eye that in time completely separates the two halves. The line of 
closure between the projection and the opposite side is indicated by 
a distinct line--the exoculata. The beginning of such a projection 
His shown in Cephaloon (FPig.6@), Epicanta (Fig./0) end many others. 


In Cistelea (Fig.Zo/) and Osmoderna (Fig.2/3) the projeetion extends 


more than half-way across the eye. In Throscus (Fig./és) the projec- 


tionnearly separates the two halves. The eyes ot Peltodytes (Fig.7 ) 
Photinus (Fig.27) and Stenus (Fie./7) are very large. Unusual forms 
lot the eyes are found in Hypophagus (Fig.%Z), wnere they sre very 


along and narrow, nearly meeting on the dorso-meson; in Cryptocephalus 


| (Fic. /06) , where they are prominent, crescent-shaped, and extend well 
ecaudad on the dorsal surface; and in Aphodius (Fig.90), where they 


fare relatively small and squere-like. The eyes of Limulodes (Fic./493) 


are transparent and almost invisible; tnose of Leptinus (Fig./a3) are 


completely wanting. 


Oculata.---The oculata is present only on the inside periphery 


of the eye as & broad ring-like sheit. It is considered of little 


importance in this study. Its general size is indicated by the dot- 


ted sres within the eyes of Cicindela (Fig.#5), Dineutes (Fig./3st, 


Passalus (Fig.2z2/), and a few others. In forms with divided eyes the 


two sides of an oculate are pressed together forming an exoculate. 


supretentorina.-- The supratentorinae represent the point of at- 


tachment on the head-capsule of the supratentorium. They are not 


thought to be primary invaginations and may probably represent no 


nore than depressions. In the Coleoptera the supratentorinae are not 


“ 


“¥atasy 24T ‘kg. It90 


E 
; 
4 
. 
wT irt 
) o/ 
i 
“4 
‘ 
at & 


/©3¥e o elqmes "agro": 9¥4a (sa eae 


he , Sota seontione. otal 


nats Seetogd ont myst wtot RE snot 


NOL 09 be 
an 


fe 


eared Se a7 sterny oe <iegs 
| ele tvedees its 

ca WRK So Fo) oo eo tee (Pe coe Me 
bfau' ott a Wa.Ol'. = roouelll et 
it a te aiedcnwowd ‘seek si eonba 
atc a 7 tas one eae” 
a (het el 


J 
i 
& 
ee 
‘ 
2 
; 
¢ 


ee ee ee sae ee 


‘ > j 2 =—" I * ? a re sos 
: , su iL REPS ae’ 


elven be a oh ee a sitobabee td sage ens) 

torte wi? 8 Rae. 
ee"q @ta 
=“geotge1t eaolivoaletae elt -a,4: 

é eiisiiahete: ef? te sicoqeotbped- 7 

1 U¢otety tem ane vey Pattie 

firdd eat ear scoryoalegy. effenl— 


ohne 


23 


prominent,.as the pretentorinae and metatentorinae often are. They 


are situated on the dorsal surface of the vertex. ‘Their presence is 


not general, occurring commonly only in the Staphylinoidea. Outside 


of this superfamily the supratentorinae are found only in Phyconomus 
| (Fic.63) and Philothermus (Fig.76). ‘The supratentoria are usually 


lattached to the inner membrane of the body-wall, but unless an actual 


itentorinae are not considered as present. In generalized insects the } 


Hiatter are generally present. 


Their presence in the Coleoptera 
should indicate a primitive state. 
Ocelli.-- In all the Coleoptera examined, no indication of any 


ocelli has been observed. 


Antennaria.-- In generalized insects there is a ring-like scle- 


rite surrounding the periphery of each antacoria. In Coleoptera this 
clerite is present, but it is distinguished from tne head-capsule by 


ridge only. 


On most heads it is considerably reduced in size, about 


all that can be seen of it externally being its projection, the anta- | 


coila, upon which the scepe of the antenna articulates. On the other 


and, in Sandalus (Fig.4&7), Derobrachis (Fig.223), Tetraopes (Figazs, 


Leptinotarsa (Fig.23/) and Anoplitis (Fig.234 ,the antennaria is quite 


prominent. The most generalized position ot the antennaria is consi- 


dered to be on the gena cephalad of the eyes, notwithstending that in 


peneralized insects the antennaria is quite frequently found distinct 
ly between the eyes. Embryology, however, shows that the antennae are 
4 


postoral in origin. Furthermore, in coleopterous lsrvae each anten- 


haria is located cephalad of a compound eye. Such a position in @ole- 


ppterous adults should denote the more generalized condition. The an- 


ennaria is very unstable in position. 


~~ 


There is hardly a superfamily 


= 
‘ 
} 
° ‘ 
by 
+ 7 


a7 
ty \ 
ra 4 4 { - 
' a 
a 
~ ’ 
f 
i 3 
: 
6 
” 
J 
7 
~ 
z 
\ 
a 5 * 
& ‘ 


nod 


a ¥t bug 82 ad? «wel? leo ne is 


4 ce eee 2 _ 2 “~- 


nfxotsrtireg 6! 6X, ate) SSN 


oughtiherss om Orie emhets 23 


sortay ttt °o booms ace 


rar Sift awe Aa? 41 OMS Be 
baott ece oanlieceer aaa 


i fo Pye) >] ‘ : Me igen 


: 
ae 
= 


eo oa eee 
uaF | : > « * > Q's ¢ @v: edz, g 


« a ome 
. : 


i rt? , Z j eee Te 


ho at. #2 ah fre 
had ek cats eye ae Aig 
: . he Sir TO orp 
Sepik) eitaes tess aso i 
dt. GUS. a2)’: Cebtwewh aa’ 
ut @ fete tony .bens (eteneg: 
age oe aegis ancy’ 
iquneth otha kt ar bce ae ead 
> oavote' severed sh on a snow 
suved qo ote eb oreo 
‘fou age <eeqnied: ag bexedane = 
coc tlevuned fot ats ‘escons saoanh atl 


= 


24 
in which it does not appear in both the generalized position and else 
where. In the Scarabaeoidea and Cerambycoidea, though the position 
of the antennaria varies within certain limits, yet it shows a char- 
acteristic location. In the former it is either on the lateral or 

ventral aspects, while in the latter it occurs only on the dorsal as- 
| pect. In no other large groups does the antennaria appear so con- 


1 stant in position. In Calopteron (Fig.Z7) and Phengodes (Fige.z7) it 


lis exceptionally large; in Dineutes (Fig/3s}] and Alobates (Fig.202) it 


is exceptionally small. 

Antacoria.-- The membrane attaching the antenna to the head-cap- 
sule is the antacoria. In removing the antennae the antecoria is 
1 often torn, and as it plays no significant part in this study no at- |] 
| tempt was made to represent it in every case. The antacoria vsries 
in size depending upon the size and shape of the scape. It is indi- 
cated in a number of figures by the stippled area: Omophron (Fig./z7), 
Neerophorus (Fig./¥), Calopteron (Fig./¥7), Phengodes (Fig.ss0), and 
} Chanliognathus (Fig J46). 

Antacava.-- The depression in the vertex, usually in the gena, 
within which the antennaria and antacoria are situated is the anta- 
eava. It is always present, so far as is known, and is developed 
into a deep socket in Dineutes (Fig.495}, Connophron (Fig.43%), Sco- 
lytus (Pig.24#], and Dendrottonus(Fie 246). 

Pretentorina.-- The points of invagination ot the pretentoria 
on the head-capsule are the pretentorinae. They sre always located 
along the epicranial suture in the Coleoptera. In generalized in- 
sects and the more generalized Coleoptera, they are situated on the 
lateral margin of the head. They are therefore represented in this 


position on the hypothetical type. ‘he pretentorinze have been 


ine vubtise bor iLademes ont diod a2 eepgeeneeeee 


raze) ent oh Chko eegose <2 rel tel OR BARS 


hes,»tc) @atafelé bar AW GSE) cetwent® ofp peers 


4 ¢hade af4e at (tea Ynesltiopie on Fryelgede “am 


i eas Vai lG » 


Z @ oe, ea 
ate = See ree 


iy 


iecy ott. M@aworlt, |, aebfonycmas9o hey seb ioee 
& Wie 225 08y orien [ ni sti8e eo tctd Ber ae i's 


,taf eft sao 2120¢fe eb 24 vondtot ee Belk 


weet =igenitetre eft teob sqQuidas ayuel 


’ wht) cebeanans tae 10 rd) rote? qoted: al “oa? 


femme = 


; } i% . ea? enitesaec sa miss dis et? 


> 


7 ay! 
sirobatna oil ,.,Sae> (r6%e ai tt ‘wenetqee” 


rl. .mqaoe i ko eqeta £04) 52% he sod 
or0 ; #42 Bet = $n.) counts ta 
, Mire eT agi ee ts Aes <?T)- xcred tole " ge <5. 
ies | er 

Welter off ab notraewqes ast ee 
tio #xe etxboatae ta4 0 Pep OE 

; owots of ea, tet of ,toegerg etewiit 
AEA2th) atueqoend> |e it) setcesie: de oe 
FL. wah amor voxnee Aas. 

igeteng ont to aobtaakeeret Ye-diaieg wil >= gard ¢ 


ayuwie oin Yodh) . etait ReTegq She OTe Bose 


titerTenes 43 motgueloe e&2 ‘ad eros. ie ie 


einotin Oth yond ei etgaeiee ene 


hafgetetdes etoteted : oa yen’ Da ARE Ot 
NAP Oo 5% ok i be fea rn roerg Ou « bens esol em 


7 J . 5 9s — 


3) 
identified in every species studied except possibly Calopteron (Fig. 
27) and Photinus (Fig.zj). In the latter they sre represented by 
depressions caudad of the eyes. Their position along the epicranial 


suture varies greatly. In widely separated groups they may be prim- 


| itively located, as illustrated in such diverse forms as Omophron 


(Fig.5), Necrophorus (Fig./3), Scaphidium (Fig.Z3), Chavliognathus 
(Fig.jo), Cephaloon (Fig.J7), Nacerda (Fig.2f), Aleus (Fig.¢9), Phy- 
fconomus (Fig.¢3), and Glycobius (Fig./o/). The position of the pre- 
| tentorina evidently cannot possess any important significance in 
every instance, yet its position may be characteristic sometimes. Int} 
the Scarabaeoidea it is never on the dorsal surface; in the Ceramby- 
coidea it is always on the dorsal surface. In this respect, the pre-| 
4 tentorinae and the antennarise behave similarly. As a matter of fact} 
they are usually associated together, but there are some striking ex- 
fceptions. The antennariae of Macrosiagon (Fig.z7g) sre located well 
ecaudad of the cephalic margin of the eyes, while the pretentorinae 
are situated at the ventro-lateral margin of the head(rig./9). 
opposite condition is found in Phenolia (fig/fg). The antennariae in 
the vast majority of cases aeopeaudad of the pretentorinse. 

The pretentorinae are the great landmarks of the head-capsule. 
On their location the determination of the presence and position of 
the epicranial suture is often dependent, and, consequently, the ho- 
mologies of large areas of the head-capsule. A case in point is that 
of the Rhynchophora,in which the pretentorinae ere located near the 
cephalic end of the snout, on the dorso-lateral margin. ‘he epicra- 
nial suture is reduced to the very short cephalic ends of the epicra- 
nial arms, and though we cannot, therefore, indicate with precision 


the cephalic limits of the vertex, its approximate limits can be 


Ayer 
ALith . 
; 3 
J asl > 
, j 
és 
' P v Yun 
La 
b 
rif 
{ 
: + 
ry 7 
& ool “ta on 
| 
he L 
7 “ 
s 
i 
«” 
x t= 
: ie : 
‘ 
’ 
‘ 
r , 
‘ 
a f 


Len 


3} 4) 
ee 
’ 
n 
4 
a 4 
j 


Ai 9 Yon? 


Z 4 
j 
; 
|= 
" 
‘ 
+ t 
‘ 
be 
+ 


s va 
wit) soretytlat yickseoy fdquxe bulfxt? stSaege eee 


tii 
ROLF feon 


ia flevbss 


= Ls 
© U ; ; 
. “> : 
, — 


- - =< ew 2 pr 


Pri 


fais P nee ne ¢ . 


, 


‘rei axn ung ba is 
ee A 


. ia) feb 


ioren 22 ae 
ia Ly) 0 a 
wos csine sie es 
4 é 
cat Ay cat 
: Sls, an te 
4, . 
f Eye ~{ 4 ra ar ; 7 
z re 2h :-t BS 


{ , 
(e ‘sioto oe 
JP 4 


: 
~? 
’ 
= 
ea 
and 
- 
$i 


a we i ae a 


ii> ALS o 


Ee <r i) ri EIA a) dal 
ey 


Ve = 


{ eae 


ey ee 


oi4 


judged, which would show the vertex to occupy nearly all the dorsal 
and lateral aspects of the snout. An exception to the ebove-mention- 
ed position of the pretentorinae is found in the Scolytidee (Bige.2#4) 
in which the pretentorinae are located in the antacava. 


Front.-- The sclerite embraced by the epicranisal arms is the 


front. In the hypothetical type (Fig./) its caudal and lateral lim- 


| its are the epicranial arms. Its cephelic limit is indistinguishable 


Since the front is fused with the postclypens. The approximate line 
of fusion is indicated by a dotted line, There is no indication in 
Jany head of a fronto-clypeal suture. The size of the front depends 
upon the position and direction of the epicranial arms. In those spe{ 
cies possessing the inverted Y type of arms, the front assumes con- 
| siderable proportions, but where the arms have been forced into es 
more or less straight line across the head, the invagination associ- 
Reta’ with the epicranial arms includes practically the entire front. 
As mentioned previously, the epicranial arms extend in this manner a- 
eross the head in a wide series of families. The front must hence as 
sume this form. In just as wide a series of families, the epierenielll 
arms are in process ot disappearing. In such cases, the caudal lim- 
its of the front can only be judged approximately. In great reduc- 
tion of the arms, as represented by Phalacrus (Fig.J2/) and Macrosia- 
gon (Fig./y), the vertex, the front and the postelypeus are indis- 
tinguishably fused into one area. 

Clypeus.-- In all species the clypeus is divided into two dis- 
tinet selerites, the postclypeus and the preclypeus. This condition 
is not present so far as I know in the more generalized insects such 
as the Orthoptera and Pllecoptiera. In some Neuroptera, however, the 


preclypeus is a large charecteristicsclerite,quite similar to what has 


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: sahidytous satigi tayot. a2 neTkeoe net eag 
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Peeve en iateerotas dat wt S aout die od awe 
(abtae efi ( \ontB) saat heokte 

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rads 
been designated as the preclypeus in the Coleoptera. It may possibly 
represent the extraordinary development and differentiation of the lag 
‘bracoria, but its size,shape and form would militate against such an 
assumption. It seems much easier to believe that this area is a true 
sclerite, and in this discussion it will be so considered. 
Postclypeus.-- The broad cephalic part of the srea between the 


fepicranial arms in the hypothetical type (Fig./) represents the post 


I clypeus. In generalized Coleoptera it is of considerable size if the 


dorssl surface has retained a generalized form. The shape, form, and 
#Ssize of the postclypeus is correlated with the position and extent of] 
the epicranial arms, which has already been discussed. In highly 
specialized forms like the Scarabaeoidea (Figs.7/73), the postelypeus 
may be even more extensive. Among the Staphylinidae, the postclypeus 
may be very lerge in Tachinus (Fig./7) end Aleochara (Fig.f7), and 
very small in Creophilus (Fig./y). In Chalcophora (Fig.z¢g) the ceph- 
alic and of the vertex is located on the ventral aspect of the head, 
and the posteclypeus is reduced to hardly more than a line. Among the] 
Cerambycoidea the postclypeus is generally very large. In Rhyncho- 
phora it is quite reduced in size, except possibly in the Scolytidae. 
Clypealia.-- The caudo-lateral projection or lobe of the post- 

eclypens is the clypealis. In Orthoptera and Plecoptera the clypealia 
is not separated from the remainder of the postclypens. In the larvag 
of Corydalis’ it.is a very prominent sclerite. The clypealia in the Co 
leoptera is often sepsrated from the postclypeus proper by a distinct 
furrow or suture. It is quite loosely attached to the postclypeus in 
the Cicindelideae (Figs.z¢uz5), the Carabidae (Figs ./2¢/29) , 

many of the Cerambycoidea (Figs.mw/os), and others. Difficulty is 


often experienced in removing the mandibles from the head without 


ei 30 


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— : 
detaching the clypezlia. The close resemblance between the Neuropterd 
and the Coleoptera in other respects would lead one to believe thet 


this similar structure in the two orders must be homologous. The pre 


Late of this furrow in the Coleoptera is wide-spread, as a glance at | 


pene figures will show. It can probably show little significance as a 
Jindication of primitiveness. It must, though, have been present in 
the primitive Coleoptera, and is hence shown in the hypothetical type 
Mandibularia.-- In most of the Orthoptera a small triangulsr areg 
is present, extending from the precoila to the cephalic end of the oc 


cipital suture. This sclerite is known as the mandibularia. No such 


Preclypeus.-- The dorsal surface of the larvae of Corydalis is 


very generalized. On this surface there is a prominent submembranous 


{ 
| 
| 
res has been located in the Coleoptera, 
| 
| 
| 
| 
| 


sclerite between the postclypeus and the labrum, the preclypeus. Such 
@ selerite, very similar in size, form, texture, and position, is pred 
jsent in Tachinus (Fig./7), Arthromacra (Fig.77), Trichodes (Fig.32Z), 
lena Glycobius (Fig./o/). This sclerite is considered the preclypewus. 
Figure 7# is a longitudinal section of the dorsal aspect of the head 
of Harpalus, and shows the characteristic position of the preclypeus. 
It is always present in the Coleoptera, though often considerably re- 
duced in size. The preclypeus is slways membranous except in Photinug 
(Fig.27), where it is chitinized and the labrum is membranous. Be- 
Sides the forms mentioned above, the preclypems is large and prominent 
in Necrophorus (Fig./3), Connophron (Fig./é6), Macratria (Fig.¢/), Phi 
othermus (Fig.7), Hippodamia (Fig.%), and many others. Very fre- 
quently the cephalic end of the postelypeus is infolded, thus carrying 
he preclypeus and the labrum with it. In such cases the preclypeus 


annot be seen from the dorsal aspect. MThepreclypeus,, no matter how 


wie 


SS dy Se 


“af Loo: 


@ 


o7 


ont BALE Sudw at neuele vote 


ah 


ag RGM syolte oMf: it, BR “ 


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é a alt | 4 i 
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mea Pos re natn 3 
- ‘ eT SCooW ¢ weds ¢ > 
a c ae i) a 4 >it Ve j j .s¢ ‘ 9 
Ves slrtaronk (of Syrah) Gey 
o ero¥ eto Ymobn Baw . (gee) ‘ade 9hOgn Lae 
« (ms 
re) 
5 t? ,#eblotel)+as MOTE LEP DOR 682 Bi 4ey 5 
PRaglortd war etend reat oly: pRAgMih aaa deta 
' & 
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en Wee comer a a ak 


29 
deeply it is infolded, is except in a few ceses, Sharply differentiat 
ed from the postclypeus and the labrum. In Glischrochilus {Fig.499)- 
end Chauliognathus (Fig./4) the postclypeus and the labrum were in 
such close approximation that the preclypeus could not be observed un 
til the two above-mentioned sclerites were Separated, and this was 
WPossible only after long Soaking in potassium hydroxide. In most of 
the Xhynchophors, due to the fusion or absence of the labrum, the pre 
elypeus could not be identified. The preclypeus, however, was promi- 
nent in Attelabus (Fig./Jo) and somewhat reduced in Thecesternus (Fig, 
W63) and Epiceerus(Fiz.3e9). 

Labrum.-- The broad prominent sclerite attached to the cephalic 
end of the clypeus in generslized insects is the labrum. In Coleop- 
tera possessing other generalized structures, the labrum is typically | 
lof the same general form. ‘the lebrum is so Shown in the hypothetical 
type. In position the hypothetical lebrum Should be, with the precly 
eus, in accordance with their condition in generalized insects, on the 
same general level with the remainder ot the dorsal surface. The gen- 
eralized form and position of the labrum is present in every super- 
fémily except the Blateroidea, Dryopoidea, Rhysodoidea, Scarabacoidea, 
pnd the Khynchophors in which the postclypeus has been infolded, thus 
Forcing the labrum onto the ventral aspect. All the superfamilies 


onteining Species with the labrum generalized, contain about as many 


Ri it in various degrees of specializetion, as to form, size, tex- 


ture and position, The labrum in Connophnron (Fig./é), Photinus (Fig. 
2%), Philothermus (Fig.70), Melanophthelmus (PFig.7¢), Eurymycter (Fig 
WS), and others, ere large and membranous. The labrum of Aphodius 

(Fig.337) is large but very thin and delicate. The labrum of Scolytus 


(Fig.%35 and Dendroctonus (Fig. #9 are probably membranous. In the 


fatdheoiwol rif Clataie’ ete Wet 1 OL SOeeRO TSR eee 


. J 7a - : 
SU. tih wwlldeoiwssa tse BF cet tens ws ee ee ke 
‘ * ‘ . - v . r ‘ S a 


ron 
ao ry 7 ge [ ara Na ba | 2 al au ’ vi aa£ Ars Ee a" 


ry av 4 ' ) i eet aey Tau? te 
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é Ahi eo 4 a ) Ss A « hy 
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latter, a significant-looking slightiy chitinized structure is located 


| 
| 


|i hecesternus (PFig.3¢7) there is a membranous area closely joined to the 


in the membrane within the mouth that may represent the labrum, In 


postelypeus that probably represents the labrum. In Hupsalis (Fig366 


jizus (Pig.3¢6), Sphenophorus (Fig.368), and Rhynchites (Fige.7%2), there 


is an area within the mouth, bounded by furrows, that may be the la- 


Hb rum. 


ithe same location and lying flat against the postclypeus a very thin 


Such are particularly suggestive in view of the fact that in 


It is present in every species, ex- 


cept possibly the rhynchophorous genera named above, where it is al- 


Iweys said to be wanting. 


Occipital foramen.--The prominent opening in the caudal part of 


the head is known as the occipitsai foramen. It is generally very 
jlarge, but in some of theliordelloidea (Figs.#6/4%) and in Connophron 


(Figs), Cupes (Fige./so7, and Rhysodes (Fig./fZ), is reduced in dimen- | 


Sions, due to the constriction of the caudal end of the head. In Ca- 
lopteron (Fig./7), Photinus (Fig./4%), Alaus (Fig./67), and Tetraopes 
(Fig.22f, the occipital foramen is extraordinarily large. 
Submentum.-- In order to understand clearly the developmental 
processes that take place on the ventral surface, it is necesssry to 


onsider a sclerite, belonging to the moutirparts, the submentum. In 


peneralized insects the submentum is not only adjacent to but is one 


of the covering parts of the occipital foramen. Such a position is 


not found in the Coleoptera. Here, it is always located cephalad of 


he occipital foramen with a distinct area between the two. In the 


no «) oxtogute herbelin 4leny tie” gees 


54). aia. of) Gatien one, (Uae Eula 


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a POeonhes at ott} ehon itt Jar Eee 

tds ai 

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sane) (rire tenes wy <t oonetel? Let tq LeGm 

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Aednen-e4 woe hich twutoee: ant ae only omar: 

iYnender ot! jet inqeneyoe eae af gisele ebro. 

rid of *omtetbe wiad Fomied iar s wi eoppeheem ad efipawind bi 

ear > dvygt .oonetel- ded petoon ea /So araag meee 


. 


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rf awd pret ondeetiah ogee swan * a 


tw ae eet eee Nl ON gh 


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Vast majority of heads this area is very wide. It is considered as 
having been present in primitive forms, and is so shown on the hypo- 
thetical type. The submentum in generalized insects is a large quad- 
rangular movable plate. Meany genera of Coleoptera show a similar 
size, form , and mobility, as in Leptinus (Fig.zZ5g9, Necrophorus (Fig. 


59), the Staphylinoidea (Figs.26j2z69, Nacerda (Fie.27/), Pyrochroe (Fig 


a 
1737), Alaus (Fig.zy), the Dryopoidea (Pics 295278) , the Dascilloidea 


} (Fig. 300), and Byrrhus (Fig.jo3). In these three characters, and the 
Jecditional one of position in respect to the parecoila, which in gen-| 
| erslizea insects is normally found beneath the submentum, that of the 


Adephaga seems to be the most primitive. The innumereble sizes and 


| forms assumed by this structure throughout the entire series of fami-] 
lilies can best be judged by glancing at the figures. It is extraordi- 
narily large in Rhysodes (Fig.3035). 
| Metatentorina.-- The invaginations on the external surface of thq 
head-capsule of the metatentoria are the metatentorinae. In the Or- 
thoptera the metatentorinee are located along the cephalo-lateral 
border of the occipital foramen as invaginations between the maxill- 
arise and the postgenae. They are not in any way associated with the 
submentum in generalized insects or in the Coleoptera. ne same rel- 
ative position of the metatentorinae is maintained in the Plecoptera. 
In a number of Coleoptera, as in Helichus (Fis.296), Stenelmis (Fiz.297 
Heterocerus (Fig.277), and even in the platystomid, Eurymycter (Fig. 
this same generalized wepition of the metatentorinae is found. Ina 
number of Coleoptera the metatentorinae are situated considerably 
cephalad of the occipital foramen. The question might be raised as 


to whether the metatentorinae that are so located could possibly be 


more generalized in position than those situated adjacent to the 


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occipital foramen. In every instance in which the metatentorinae are 
located cephalad of the occipital foramen, a suture connects the met- 
atentorinae with the foramen. In only a few cases does the suture ex 
tend much farther cephalad than the metatentorinae. This suture in 
the vast majority of heads studied does not extend cephalad. It is 
readily seen how the metatentorinae might be drawn cephalad and as a 
result a suture be formed marking their line of migration. In such a 
process one would naturally not expect to find a suture located ceph- 
alad of the metatentorinae, and in the cases in which the suture does 
extend so it is easy to understand that the force of the cephalic 
pull might have been communicated to this region, producing in conse- 


quence a suture or invagination. Due to the shape of the head, it is 


there would be formed a suture cephalad of the metetentorinae, mark- 
ing the line of migration. In this discussion the generalized posi- 
tion of the metatentorinse will be considered as that of their general 
ized position in more primitive insects, at or near the occipital for 
amen. 

In the development of the coleopterous head the metatentorinae 
have shown a tendency to migrate cephaled. The cephalic migration of 
of the metatentorinae and the ventral migration of tne pretentorinae 
and other structures, were no doubt due to the same force, the result 
being a closer approximation of parts,which naturally supplied in- 


creased firmness to the head's mechanics of operation. The Dryopoides 


Show the most generalized position of the metatentorinae. Genera of 


this superfamily nave been mentioned above. The Elateroidea 


probably possess the next most generalized metatentorinae, such as in 


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Sandalus (Fig.287) and Alsus (Fig.Z#). The Cucujoidea show the meta- 
tentorinae just a little removed from the occipital foramen, as in 
Megalodacne (Fig.3jz), Anchicera (Fig.¥/4), Philothermus (Fig.j/7) , and 
In the geners of other superfamilies, as Nosodendron (Pig 30g 
noplitis (Fig.3s), and Tetraopes (Fig348), almost the same degree of 


rimitiveness is shown. The large superfemily Mordelloidea show the 


ent through the group. In the Bostrichoidea, the Scarabaeoidea, and 


he Cerambycoidea, the metetentorinae show considerable variation in 


position. This is shown by a comparison of their position in Bostri- 


ehus (Fig.333 and Sphindus (Fig.335.. In the families to which Tetra-| 
pone (Fig.74f, Calosoma (Fig.zs), Cybister (Fig.75f, Dineutes (Fis zosr 
| Necrophorus (Fig.zs9), and Glicobius (Fie.3¢7) belong, the metatentori-| 
nae have advanced very far céphalad; near to.the submentum. In Pho- 
tinus (Fig.z7z) and Chaulioenathus (Fig.z7¢) they are located on or 
quite near the psracoila. 

Maxillaria.-- There is a nerrow plate surrounding the lateral and 
caudal margins of the occipital foramen in some generalized insects, 
between which and the postgenae the metatentorinsae are invaginated. 
This plate has disappeared in the Coleoptera. 

Cervix.-- The structure connecting the head-capsule with the pro 
thorax is called the cervix. It is normally composed of membrane, 
and a number of cervical sclerites. The size of the cervix depends 
upon the size of the occipital foramen, and the degree of mobility of 
the head. In the Lampyroides and some other forms the cervix is very 
large. ‘The cervix in Rhysodes (Fiz.305) is composed of tough fibrous 


membrane, quite different in structure from the normal cervix. In 


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34 
nearly all of the Rhynchophora the cervix is heavily supported by 
strong tendons attached at its cephalic end. ‘hese tendons take cere 
oft the added strain on the cervix due to the elongation of the snout.| 
There are more or less small cervical tendons appearing occasionally 
] throughout the whole series of families studied. In Molamba (Fiz.2Zé6/) 
4 the cervix is invaginated within the prothorax, doubling upon itself. 
7 The cervix in Bostrichus (Fig.jg33 doubles beck upon the head-capsule, 
qwhich is produced into a round projection. 


Cervical sclerites.-- The most prominent and persistent cervical 


sclerites are the pleural cervical sclerites, the cervepisternum and 


the cervepimeron. The former is usualiy the larger ot the two, and 
articulates at its cephalic end either against an odontoidea or sim- 
ply egainst the unditferentieted area surrounded by the occipital 
foramen. The latter usually extends in. a different direction from 
the former, and articulates st its cephalic end with the cervepister-| 
}num and at its caudal end with the prothorax. In Cantharis (Fig.2775/, 
| Macratrie (Fizc.733), Psephenus (Fig.Z”7, and others there is a single 
large selerite present. In the Adephage, in Leptinus (Figc.2o5y, Hy- 
porphagus (Fig.329, Phyconumus (Fiz.707, Pseudocistela (Fig.32Zg, and 
many others, there is a Single small subcirculsr sclerite present. 
Both of these types probably represent the cervepisternum. In a very 
large number of genera cervical sclerites are always wanting. They 
are poorly or not at all developed in the Cerambycoidea. None of the 
Rhynchophora studied possess a cervical sclerite except Hurymycter 
(Pig.3/), in which it is very smali. The ventral cervical sclerite 
is the cervisternum. Nacerda (Pig.23/), Cucujus (Fig.39/#), Hister (Fig 
Z7o), Tenebrio (Fig.3%) and a few others, possess two small small cer- 


victerna, while Tomoxia (Fiz.28% possesses a long narrow one. The 


Ms 


i “cc be2 toggua  Yslvned Br viveies wil che 


Betas otat anopasy eed? ehBe Cilarged OFF hg 


: ese ort ro no ltonsoLe BRR er ab BLP Tas: Hs"? Gy 
Dytiondtheoss gatuecgge Bbbbnst novi ha 

} — ' E 

|) a fa) Adele el . .begises x an a 1s 8 '@ aOLieg: a 


ecoay niiieacos estacsidta. see sith eae 


: —— ‘ - : cy peak 
aeelenges-r aad cig tox { Ls ee Oy Ge) j ae 
, A ¥ 
swetiperurs on SS 
4 
$ A F. 
ZAC 4 fain tog ia Fat 4 -rsBoOPee 
\ : ‘ 
baa m telqevrss 2 ivice (ssa 
= 
4 * s 
; Pics ox Th Pe o 2 ; 2. %2Sac8 


oe 


cl .4“entetalqeries. sr--3 jel Lil's ut 
iijasw nyatia sts vet) aie2 izat 468 ire 

s?! fd eabtor inks mit! F P hashs ¥ . a 
so‘ srowyire fives® of tcelie 246 ivaae 6h wade 

- alee Laolivied. Latiget.os b itaon rity &F ine 
29) cofati. ,(We.7i4) GB. buds as he usd) 2 eo, ae 


eraeo Lican itans .ow? 2898209, ateddor wete het 


af? .one gettag. seek a -peeenemag s.s2. Lxoma?.-s 


o ; ee we es rey 
é 


35 


ecervisternum is on the whole of infrequent occurrence. ‘The dorsal 


cervical sclerite is called the cervinotum. It occurs even less tre-| 


quently than the cervisternum. Hydrous (Fig.//) possesses a subquad- 


rangular distinctly chitinized cervinotum. In Aleochara (Fig./S) the 


eervinotum is divided into two distinct lightly chitinized subtrian- 


gular sclerites. Two much larger square-like sclerites are situated 


eaudad of these. All ot the Hydrophilidae and Scarabaeidse possess 


 setaceous caudo-lateral sclerites, which do not seem to be present in 


] other’ genéra studied. 
Odontoidea.-- Owing to the strong chitinizsetion of the head and 

1 the close fit of the head in the prothorax, there is little need for | 

#a special process or projection on the head for the articulation of 


ithe cervepisternum. Such a process is called an odontoides. The few 


linstances in which it does occur are in Scaphidium (Fig.37), Eucine- 


}tus (Fig.7y), Cyphon (Fig.gzg, Byrrhus (Fig.gzy), Nosodendron. (Fig.¢zs} 


Phyconomus (Fig.g3o) and Arthromacra (Fig.m. The latter genus, it 
is interesting to note, possesses no cervepisternum. 


Postgena.-- That part of the head-capsule on the caudal surface, 


W]mesad of the occipital suture and ventrad of the occiput, in general- 


ized iorthopterous insects is the postgena. The occiput is considered 
as extending to near the middle of the dorso-ventral length of the 


occipital foramen, thus limiting the dorsal extent of the postgenae. 


The latter sre widely separated from each other by the occipital for- 


amen, ‘he lateral parts of the large submentum distinctly cover the 


mesal parts of the postgenae. In the Coleoptere the occipital suture 


separates this region from the remainder of the head-capsule, as in 


generalized insects, and the occiput is also indistinguishably fused 


with the postgenae. But the postgenae, instead of being widely sep- 


e\selt) ex@apoe la 22> -.esvisoerios Fee 


- f ae" ye MW rye » 
mat : a de et oleae 
Lee er a ae ee eee yt 


axob 288 Veeoeeiund Xedos etrat he kee pee 


is neve gteebe ID) FR ga ives a i 
i 4 yy? EBON WES} tut hy  mipenedg 


haan: eho. . ; (Ocul s Ost sau Sf fy ge vis 


“ge PP of coe PoP ot ; 2 > "0 ee ntest ent 


mbbabiretes Ks efi a a seeks 

vo MAU BEY rtihideesd ax ory royal 
ion | abo hatin vet Ze a Re D kia . 

> usin Sele ua) 
olan (On pecaeemigm 7 safe. 

Liki: eivaqe? * eed alt hy vine ut 
“oe , ~ bestasyl hee “expat 

; | “bis -Ansateay oa ee aoveen) 
ius 80nd ey Soi aaa at * 


af ygaene-faot ent’ Lo reba laa an? om 


~ 


a 


to bine Peer ler: 4a ie e ty hoeo ado’ 
febiw “aioe ta baw Mad ” spuiegrien, ath 


36 


arated and their mesal parts being covered by the lateral parts of 
the submentum are directly connected by the broad area loceted be- 
tween the occipital foramen and the submentum (Fig./ ); and the sub- 
mentum is attached to the mesal portion of the cephalic margin of the 
| postgense. This area is indistinguishably fused with and undifferen- 
tiated from the postgenae in the hypothetical and the more general-' 
ized types of ventral surface. 

In generalized insects the submentum is joined to the cervix by 
the undifferentiated part of the cervix, the cervicoria. In the ceph} 
alic migration of the submentum, a part or all of the cervicoria must 
have been drawn in behind. ‘The postgense alone bordered the lateral | 
margins of the submentum and the cervicoria. A possible line of de- 


velopment may have been the chitinization of the cervicoria and the 


ee of the sutures separating it from the postgenae. The 


area between the occipitel foramen end the submentum would in 
J such @ ease be a modification of part or all of the cervicoria. It 
jis much easier to believe, instead, that because of the membranous 
condition of the cervicoria, no strong resistance was offered to an 
approach of the mesal margins of the chitinous postgenae, which would 
migrate thus to serve as a firm foundational support for the svubmen- 
tum, as well as to strengthen the solidity of the head-capsule as a 
whole. The mesal margins would in time approach so close as to press 
the two sides of the cervicoria into a line. Fusion of the mesal mar 
gins of the postgenae would finally take place, and the cervicoria 
would be divided into two parts. One part would be attached to the 
submentum entirely separsted from that portion of the cervicoria bor- 


dering the occipitsl foramen, and would be finally reduced to a su- 


ture. The other part of the cervicoria would remain as a portion of 


be ne “§ ¢ 48 winaes nino ePeayiyeeuren? far 


‘Gra ott tol 7A eft) aera ot? (bre RAR 


3 ‘o tear OF Loy » Oe 34 I (foe, 2eaae aie wey 


tc “ixee Laséhel atc ee Meuvics arcee eek 
od beteng? ahem bacté Beh gv taloe tote 


ey Phue fine aleds ay i Or, Pan 
ie b rj é ra { } ’ y F SALE) 
: 145 


sub 
- 
Ar f 7 fy : Hi fad Ae 
b + u 
my Cask 4 “4 uve 


2  » etre 7S ; LSE": 364, ep 


~ 
— 
aa 
2. 
I 
re 


ae ey est “fie lt Rowdy obey 


¢ etieniwves et? ty nodded lam gether heen a 
ube 
ee fk sata el 


‘ i, i mn a ae 
Pe i aa «+ =n car pena i —: — P Pg 


enna 


o 


ut wo OF beagene ~tiwl? «> thew fee on 


oO” 
the cervix. Whether the messl margins of the postgenae would ap- 
proach the meson as broad surfaces, or aS narrow ones and later elon- 
gate,it would be difficult to state with certainty, though the latter 
would probably happen. In the primitive coleopterous head it is not 
unlikely thet the head was much shorter, and that the area between 


the occipital foramen and the submentum was correspondingly shorter, 


iso that there would not be the necessity for a broad fusion of the 


ymesal margin of the postgenae. So far as observed there has been re- 
aval in no coleopterous head any marks of the line of fusion of the 
} postgenae. 

Abundant evidence is found for the conclusion that the broad a- 
rea between the occipital foramen and the submentum must be a part 
of the postgenae. A similar condition as in the Coleoptera is pre- 
sent in many families of insects. In many Hymenoptera the messl mar-| 
f gins of the postgenae are fused between the occipital foramen and the 
larticulation of the labium. That this is the true interpretation of 

7 this structure is shown by the fact that there is often a suture on 
the meson showing the lime of fusion, and is further substantiated byt 
the fact that this area bears on its ventral margin the paracoilae. 
In all cyclorrhaphous and orthorrha phous Diptera the area ventrad of : 
the occipital foramen is a continuous chitinized piece, similar to 
that found in the Coleoptera, and is derived from the fusion of the 
mesal margins of the postgenae. 

It may be well to add here that the expression “cephalic migra- 
tion" as applied to the changed location of. such a’ structure as the 
submentum, may be only relatively correct in its suggestion. Very 
probably the occipital foramen in Coleoptera has migrated dorsad, at 


least it has sssumed this typical position. Such a migration in 


y e Tad 
4 
, 

a Ie] rx | i 
; ? 
» YO; 
i ted! ak 


fae Hag 
, "tarry 
' 
4 Ll By 
) 
or 


‘ie omewefan) wait Se sateyor Enlai, ce ae 


vrate x! t ab ad ivitnios fet ble 


AA2F SH Pete ier Pest] «0 Gia ieee 


‘ag ; nT oe J ea eS 7) i ‘ aye) we ee 


7 + : 'y | thw 
a 4 7 
Seen A eee ee ee wh hese ot - ae at om = retin. 


Few tune wittstiiwe is ,oouerieg, even) 
dh wtekeeteuritirie «Fats eieg 


ner dtouerqselds  ovicioitg sm eee 


wee fh eve Wet mruonde& Sir; oe 


fae | mosis eb tack ‘booe? 


oe ot «eee 


- se ceaigal 80M oy 
uises* petoseuund ai’ todd ate oe as 
guadt: «= deuiet¥o uel TER or Sr elt al 
‘Tteeguie ell ot tt a 


tet ao 6a 2a goesor rds sage tO} | 


in » deve  igobeiedé Leatny? 
° 20s ost ee ST =i 


38 


itself would produce an elongation of the ventral surface, with a con 


sequent production of the broad area between the occipitel foramen 


and the submentum. Whether the force exerted on the head was direct- 


ed more towards the ecephalization of the submentum or towards the dor 


salization of the occipital foramen it would be hard to say. ‘tne con 


dition resulting would be in either case approximately the same. For 
i purposes of convenience and simplicity of language, the expression 
i"cephalic migration" is used in this discussion. 


This broad area between the occipital foramen and the submentum 


is sometimes designated as the gula. Usually, though, the term gula 


is used to indicate a sclerite, on the meson of the ventral aspect, 


separated from the remainder of the head-capsule by distinct subpara-} 


llelsutures, the gular sutures. The term gula cannot satisfactorily 


tbe appoied to both an indefinite region and a distinct seclerite. The 


sula is considered in the latter sense in this discussion. For the 


present the area between the occipital foramen and the submentum will 
} simply be designated as the mesal parts of the postgenae. 


The cephalic end of a postgena is usually modified into a rather 


iilat area sloping towards the meson, upon which are situated a para- 


coila end a posteoila. This flat area is distinguished from the re- 
mainder of the postgena by a distinct ridge, which usually extends in 


& deep curve from near the lateral margin of a postcoila to near the 


caudo-latersl margin of the submentum. Sometimes this ridge is di- 


rected towards the occipital foramen, as in Leptinus (Fiz.253), Can- 


tharis (Fig.279), Macrosiagon (Fig.7Z%3), Tharops (Fieg.79z), Psephenus 


(Pig.z75), Helichus (Fig.2%), and others, Attention has already been 


ealled to the fact that the cephalic portion of this ridge represents 


the cephalic end of the occipital suture. This modified part is 


~ el Ue oe | 7 : z 
' Me nabs ee Way 
\ f ‘ 10h» eames & af ow inte 4~ 4 1 pp hed > + es mpi = rye . 
pier A OS a a aa el Wats Stal elle AY wha ! ere 
; 
ree 


R anetoat (ue ee iyeg Sih qaayeed 15 


cow fet etd: nocteicaxe 

if n : J — 

meh ect gbaamoe es main meyE edi 

4 F & . : Ps y if 5 

| of dies ed Picow 72 meageg Lee ap e 

| ah inan oat yieteminongce vier pitie aba 

‘ : is ; 

; ‘ a d 

i antesagunze eff. .4A2 tj PLODLaMsAe aah 88 
De ee 

Garr? peer Gia 0 how @ ols: ; 2 ickt . Ke eo 


: ,  NEMOAL ) -2iimm ee an, hePates 


' é fae mee ¥ 
me | yeriretlor 3 vet pine Lay otsorteiih ae 


ct i 


. hedtveot a offer be) i naews ase [vie rh eog a 2 bs) mA <a 
ewe @ Pere - low Ar go, beeem Ch” eieiaaed aie 
: . ; ees 


an 


il i Vt copeboenies. ‘a ~~ inten @fiite, Te 1gaa 
«cad. . (eet. .t@) oontiaet ef Be \remesd Hed L420 9p 3 7m 


| ir) 
' OL glee es Sates PBA. aTy 


a r 1% oo 
Powe! ' ian coobingiia ical? 6 pee, . men Ee 
: + ; + 


> Ls Pal 2 
Bria. 1 mbit aid? tf celta aan 
' : ; : r a Me 
f /l tury butifow hii! vem aoe ae 
‘ , Z 


a il 
st ow nn Py a ee t 


ay in al» 7 as \ 


39 
reduced in size or indistinguishably fused with the remainder of the 
head, in Photinus (Fig.z72), Eucinetus (Fig.300), Alobates (Fig.3z5), 

1 Tenebrio (Fig.3z), Hyporphesus (Fig.3z7), most of the Scarabaeoidea, 

Sphenophorus (Fig.36%), Hupsalis (Fig.3¢60), and Lixus (Fic.3¢¢). 
Gula.-- There seems to be considerable confusion as to what 

stitutes the gula. Some investigators consider it as simply the 


4 tral portion of the ventral surface of the head-capsule. Others 


f strict it to the distinct sclerite appearing on the meson of the 


i tral surface of certain insects, including the Coleoptera. Since 
this sclerite is distinctive and characteristic it needs a name. The 
‘term gula is quite satisfactory to apply to it. The origin of the 

| gula also rests in considerable confusion. Comstock and Kochi (1902) 
| consider it the sternum of the cervical segment, and to strengthen 

| their argument they cite the situation in Corydalis, in which they 
state that the sternellum of the cervical segment is retained back of 
a the gula, and they figure it as the small quadrangular sclerite imme- 
| diately caudad of the gula. They do not mention the very much larger 
rectangular cervical sclerite caudad of their "sternellum". The lar- 
ger sclerite is suggestive of the sternellum, and the smaller one 
would then represent ee "sternum". It is generally stated that 
the gula is derived Soon lavienat sation of the neck membrane or cervix 
In the discussion preceding, dealing with the metatentorinse and the 
postgense, I haveattempted to show the impossibility of such a deri- 
vation. In many orders of insects structures on the caudal aspect of 
the head have been designated as the gula. My interpretation would 
prevent the recognition of the presence of the gula in any insects 
other than the Coleoptera, Neuroptera, Trichoptera, and possibly the 


Mecoptera. 


Gwened was, lo tebe « (30.5 7) 
1 > ? 


an * wr) Pei. ale ae dee react qitetstoee es 


i 
Sp oon 
: ; * 
ey 
He 
| ae , 
7 
‘ ; { 
al) on “> 3 re 
4 
~—_ » 
-_ ¥ 
Sle 6G 
- | ‘ 
es it f 
7 ie } ; "> fi 
j “a [ “a en 2) ° 
j 


ee 
i — 
; 

: 
1 

4 ; 


TC 
f 
ry 
‘ 
4 
f 


¥ 


~ 


Sites #4! bo. seth bow wD edosen sto 


= 


ostedéera , A994) soret ogl, ee 


“® 


eae ft ripe 4 >i POPE 
f 3 7 : f Ow 
. weG -s ; 
3 fit is 
’ e 
é : ’ Het 
+ ro a ie A 
pe rf i} 
f z ‘te 1 
; ‘ és 


" prc to “De hraw wr ivetae® 


EDL Sater e weal le Ge 3 hve § | 

; el 

126 f. tate” Be nadia 

yey = ae Gti ives Ald oe anh 
soineote. an?’ -i¢ ie. soklieak ase 


if ielmamiml add qwode: of Belg 


' 
a 
ot 


[titel <M ip Sk ne Loteaste 
ol eteq wft to-wanepete «iio Eoraes 


bhisy 8 164 certaget iste quved, 


40 


= 


practically 
In some Coleoptera there is,no gula (Figs.267277 ), whereas in 


others there is a complete gula (Figs. 49/ 25¢6 ). Between these ex- 
treme types there are found in the Coleoptera all gradations of a gu 
la in process of formation (Figs.29/ 256,333). Such a range of vari- 
ation in a structure simply emphasizes the importance of a compara- 
tive study of a large series of forms. The question as to which is 
the more generalized condition, the presence of a complete gula or 


i the absence of one, has been discussed previously, with the verdict 


in favor of the latter condition. It would then be impossible,othe 


f reasons not being considered, for the gula to be the sternum of the 
cervical segment. The gula is the sclerite formed by the migration 

} of the metatentorinae from the occipital foramen towards the submen- 
} tum, and must be derived from the postgenae. In this migretion of 

i the metatentorinae sutures are produced which are known as the gular 
sutures. The area between the gular sutures is the gula. It is evi- 
adent, then, that the presence end extent of the gula depend upon the 
position of the metatentorinae and the sutures which they produce. 

i The gula is complete in all the Adephaga except Peltodytes (Fig.253), 
in the Silphidae, the Staphylinidse, the Sphseriidae, the first four 
families of the Cantharoidea, and the Scarabaeidee, except Pseudolu- 
eanus .(Fig.343). Isolated examples in other families of a complete 
gula are Scaphidium (Fig.26yn, Helichus (Fiz.Zw%), Cyphon (Fig.3u), Byr 
rhus (Fig.3o3), and Leptinotarsa (Fig.354. The gular sutures nearly 
meet in Gastrolobium (Fig.z63) and Creophilus (Fig.géq, and are entire 
ly confluent in Necrophorus (Fig.Zs}, Scaphidium (Fig.269), Hister 
(Fig.270), Phengodes (Fig.z73), Chauliognathus (Fig.z%), and in all the 
Rhynchophora except Kurymycter (Fig .36/) , in which genus the gular su- 


tures are very short and widely separated. The gula in cases of con- 


a~feo lo andere nh slim wit .easeged ec 


mo 40649 cere Ag \ wes Nth 3. regs) olay oat 


Seetide cit whiawet wematot [x tetong Seeoeeenn 


rata: Lf) 28) 2S é es ae ae a 


pi kensyiente 
|) NOS 504 eR el a Pt all 


* As 


os) o 06,88 ham Jie ae noeret af) at & pula 
al t ¥ } cine 5 5 196 ‘ ei ts & Sell + \¥E Pa % , £y tag el ; 
-aragcov » tc sowetical ect wee aices: SOR 


ef on * ~ te nefieuge eT ~umProl 7th 1 hese lee 


16 ALLS a eee . eat ie ane nob) ihe 


B; pid 44. | Pe Bi) Aes ¢ tio. , be othe 
Soy asash wir Oe vemo: bins({ae .Got)/an oa G oi vs 
; ae 


noizgesain aia? 1 (Gaus )foq sue eee 


wid. yi 7 Casa ‘ ‘ { a na on aati a 
naut¢ fact Hi va ers ald z in aa te 


+ 


“tees Lo) BENS Ls me PEP ae F CBy ae ak ‘sted 
earer* ts rt Ae os be Enua ele’. S Af oie, x 


on. eotdiue, sein ets Ree 1S ee ats sitqesth ae 
‘ aay Ae > 
eliieoutes £0: arene Sapo oe ia 
: we ; 
bio, was. ott) aed) iveeod eRe (%) ‘sere dave y 
a ct . 

if. ot J08 Se oD parcfense i Iaetl Sa 

me Seiad 


1; i= ej? anaes in idw (iE sf ea) a 


“a 


H 5 : tsi , ea. 94 \ 


pane 


fluency of the middle portion of the gular sutures on the meson is 


41 


simply invaginated. The gula is complete but extremely short in Cal 


opteron (Fig.z7) and Photinus (Fig.z7z). The cephslic ends of the gu- 
lar sutures meet on the meson near the occipital foramen in Philothe 
} mus (Fig.3/7) and Anoplitis (Fig.357). The gula is partially membran- 

ous in Nosodendron (Fig.309), Sitodreps (Fig.33z) and Plesiocis (Piz.334 
) Gart.of-it is deeply invaginated transversely in Scaphidium (Fig.29), 
# Chauliognathus (Fig./7), Helichus (Fig.29¢), Stenelmis (Fig.297, and 
4 others. 
Epicranium.-- Most of the sclerites of the hesad-capsule are so 


closely united in some Coleoptera as to appesr as a single piece. 


| Straus-Durekheim (1828) so regarded this region in the head-capsule 


| of the Miay-beetle, and named it the epicranium. Used in the sense of 


| indicating a closely united area, the term seems satisfactory and may 


often prove convenient. It can then be only a relative term, includ- 


ing at times no more than the paired sclerites of the head, when the 


epicranial suture is well developed. Occasionally, the preclypeus 


and the labrum are closely joined to the head-capsule. In such cases 


the epicranium would embrace these structures, also. 


Paracoila.-- The small rather concave projections at the caudo- 
mesal maxeins of the postgenae against which the maxillse articulate 
are the psracoilae. Piney gant usually easily identified,.as in Molam- 
ba (Fig.z6/), Pytho (Fig.z@%), Heterocerus (Figwz9y, and Phymaphora 
(Fic.wy7). They are usually chitinized, but are membranous in a con- 
siderable number of genera. ‘The paracoilae are insignificant in size 


in Gastrolobium (Fig.Zé3), all of the Hlateroidea, Cistela (Fiz.32zy), 


and Tenebrio (Fig.326). In generalized insects the paracoilse are sit 


uated beneath the submentum. They are found in a similer position in 


— 
| 
: 
, = 
a 
ic 
| 


eoge|c 


ait op eti tafe od? to nO RP aera 


ix planers fa¢ bialqrer el ki wt 4 


thks Ghisrtn dh SEE» « iaxk. 528) somtt ond i 


. 
ag 
1 
* 
aff 
é Ae 
oqgs a¢ 
r i , 
’ . 
/ iis 


? ‘ = 
+ doewenr LAT eefoo say [ieee ooRee hd os 


oor 34 aise ect «Sten )) eRe 


’ 
rr 
b 
a 
' 
y 
> 
b 
” 
& 
5 
> 


i 
net rilzsaw at? toltw.te: + CA SRE 
<i oe, @otritaehl. plies Viler cs ats Yas 
I 


sovits Spa , QRWAEE) Betacotsre: .. wk APS 


é 


caeldting lanl ote oeiioogisd sc. dieehe: 


quoratdaem ot@ ta¢°*, Geainid tao Cl nna 


pioerar. mob tovatea SE edd- 20 iie .; V ee 
pai 


‘casera sat utaseei. “bamntereais et iv 
pain 


yafi¢ia ah) Seocd ete gaa aut 


=) 4." a ee eee 
e baal 7 


42 
the Adephagea (Figs..1#725/), in Hydrous (Fiz.zs¢), and a few others. 
They are also always located on the postgenae in generalized insects. 

j This position is of great importance in determining the identity of 
the postgense in specialized insects. 
Postcoila.-- At the cephalic end of a postgena is always found a 
diosmnet crescent-shaped acetabulum-- the postcoila, against which 
y the postartis of the mandible articulates. Throughout the genera the 
| posteoilae exhibit some degree of difference in exact position upon 
the postgenae, in size, and in degree of shallowness. Their form and 
position are indicated in Calosoma (Fig.250), Spheerius (Fig.7693), No- | 
toxus (Fig.7%), and Derobrachus (Fig.3¢). In generalized insects the 
postcoila are also always located on the postgenae, and their posi- 
tion is of equal importance with that of the pareacoilee in determin- 
ing the identity of the postgenae. 
Precoila.-- The rounded condyle on the ventral surface of each 
4 clypealia, against which the preartis of the mandible articulates is 
1 the precoila. It is usually crescent-shaped, but sometimes, as in 
Peltodytes (Fiz.253), Cybister (Fig.25¢), Creophilus (Fig.2é9), Endomy- 
ehus (Fig.3z0), Dichelonyx  (Fig.33x), and others, the form is spher- 


ical. The precoilae are large in Macrasiagon (Fig.7f3), Epicauta (Fi; 


| 254), Tharops (Fig.z7z), Chauliognathus (Fig.774), Bostrichus (Fig.333), 


and others, and small in Dineutes (FigwZzss}, Photinus (Fig.27a, Heli- 
ehus (Fig.29%), Derodontus (Fig.3/3),and particularly so in Psephenus 
(Fig.27f). The precoilae can readily be recognized by their character 
istic form. Since they are always located on the caudo-lateral lobe 
of the postclypeus, or the clypealia of certain insects, they are a 
great aid in determining the limits of the postclypeus. 


Tentorium.-- The endoskeleton of the head is known as the ten- 


a 


am 


if 


i 
o 
A 
‘ 
+ 
‘ 
¥ 
a 
. % 
i j ) 
< 
wi 
4 
’ 
Bal 
pa] 
; o 


‘Reis h) aueshe! he Pina. 


“lt BeGatfece ad. we has tortie = hy 


TL iMiie Si 


iy 
Peri 
be iid 
. 
Lf 
js 
‘ 
‘ 
+ 
f ; 
‘ 
+y 
a 
) 
“> 
. > : 
- ' 
| yer 
t 
e T 


a 

2h 
tJ —~— 
<u 


4 
ts 
ss 


aod ult De doteteteall 


a 


' 33 8 4 bnew 
a 
i226 etd see oer 
sal ‘an Ci ty [Cece Wy 


if, = 
od ue: = | ° 


(Neh. 3 i mrs Ss) a % * 


sia? fig 


‘ = ' 9 Pi 
a7, y 7 , 
ivavpnecl Lapis <2 
+ AZ he 4 


44 
rp 
rs) 


yee ie? od oe 


43 
torium. It is composed of two anterior and two posterior arms or pro 
jections, and their various modifications. The anterior arms arise 
from the pretentorinee, and are known as the pretentoria. The pos- 
terior arms arise from the metatentorinae, and are known as the meta 
tentoria. In generalized insects one end of the pretentoria expand 
] along and are continuous with the epicranial suture. iach extends 
in a caudo-mesal direction, and narrows for a short distance, then 


expands along its mesel margin until a fusion is formed with the pre- 


| tentorium of the other side, producing the cephalic bridge or lami- 


7 natentorium. The pretentoria separate and then fuse again farther 
cauded with each metatentorium. The metatentoria extend cephalo-me- 
sad a very short distance, their mesal mergins expand and completely 
| fuse on the meson, producing the caudal bridge, or corpotentorium. 
The fusion of the pretentoria and the metatentoria is supposed to 
take place along the cephalic margin ot the corpotentorium. The dor 
sel projections arising from the lateral margins of the pretentoria 

f and extending toward and attached to the dorsal well of the head, ar 
the ee achteric. The ring-like plate surrounding the inside peri- |] 
iH phery of the occipital foramen is indistinguishably fused with and i 
& part of the metatentoria. The tentoria, as a whole, are distinctl 
ehitinized and well-developed. The typical condition of the tentor- 
ium in generalized insects is practically duplicated among the Coleop 
tera. The hypothetical type (Fig.370) has been constructed with this 
Similarity in mind. The greatest difficulty experienced was in de- 
ciding upon the primitive type of laminatentorium, whether it should 
be represented as complete or incomplete, that is, whether the two 
Sider of the laminatentorium fuse on the meson or not. Many Coleop- 


tera that in other respects are quite generalized do not show a com- 


3 P 5 4 
4 p 
may iO. anita etree cmt dees LotreP ae oard’ te 


wtiud dee tetaet an od. 3) eee Eyer. “aioe 


we fy nnAs «il ahera - ae: eae = i tewOtcs, tis bev 


__. ¢ } ~ 
Rha ed] weg é Say TVR LTO 
itt iAs 7 yp) aie = hp © a P is y obi 


SEMIN Nhe? rata Jath Eolas 


Soci fae" fae = nae Ts 7 }® Wot 
meron OLEIg¢e90 ansvaa wl { ) ai 


. ! ~ 4 
i Pesaougas & i : i iq 
et | PE 2 eels Ov i : ee S251 


'g 
y a | ,ii6 E | ad a é ‘ft 
m = 
‘ lp 2 & + _ * 
ms J Pe iy. t 3 2 \ a 
+ t I | 
Ee Bet Aina! ais { ; t 


ee od aoa Derea tic tact 


al ; iW Preoreigagee yr (pol. ii 
i wenw we Pindsean keel 
yo! , 2) di, OT ; #} ir Ae 2 4 of f5 EGE? 


mot C1 ZA fo 338 owen sAt ee aoe 


nen # FONG “oo Of Dene Dip aeg ee ng ae 


dems oo 1 ney 6s eee iam. dibeieaiiass : 
: / 


44 
plete laminatentorium, as Tetracha (Fig.37/), Omophron (Fig.375), Di- 
neutes (Fig.373), Leptinus (Fig.ge/), Tachinus (Fig.337), and Stenelmis| 
(Fig.7/g). The hypothetical laminatentorium is represented as nearly} 
meeting on the meson. At least, such a condition is thought to be 
not far removed from that which actually existed in the primitive Co 
leoptera. lLimulodes (Fig.339), Burystethis (Fig.¢0¢), Pytho (Fig.4o7), 
Philothermus (Fig.#7), Melanophthalma (Fig.#39), Hyporphagus (Fig.gse 
Sehinded (Fig.7%), and others, possess practically no trace of a lam 
inatentorium, but well-developed supratentoria. The latter sre fair 
ly wéll developed in Photinus (Fig.37g), Collops (Fig.s77), and Alaus 
(Fig.g7z), but there is neither a laminatentorium nor a corpotentor- 
ium present. There is no trace of any one of the three above-men- 
tioned structures in Tharops (Fig.¢3)and Throscus (Fig.wy). Phelac-| 
rus (Fig.wz and Eurymycter (Fig.vs3) possess only the rudiments of 
the pretentoria and metatentoria. The tentoria of the Cerambycoidea| 
(Figs. 474.4779 ) are very delicate and membranous. The pretentoria, 
the metatentoria, and the corpotentoria are always complete, while 
the laminatentorium and supratentoria are always either rudimentary 
or absent except in Pachymerus (Fig.v), in which the supretentoria 
are present. Other genera showing completely membranous tentoria ard 
jnumerous. Among these are Cupes (Fig.vo), Buryste thus (Fig.¢oc), Rhj 
sodes (Fig.¢zé), Languria (Fig.#2), and Pseudocistela(Fig.ws). 

Pretentorium.-- A generalized condition of the pretentorium, 


|characterized by a strong chitinization as a whole, and possessing a 


joroad flaring cephalic end, is found in the Adephaga (Figs.374 377), 


l 


|Leptinus (Fig.3%#), Necrophorus (Fig.3%4), the Staphylinidee (Figs.3%9] 


386), Georyssus (Fig.g26), Hucinetus (Fig.g2/), Dermestes (Fie. $23), 


and Derodontus (Fig.494). There are all degrees of gradation present 


a 
pt 
. * : i) 
o Sh , (owe ~] 
‘al yi J *< 
ner reat Ss 
Oe r ba 
. 
& * 
a* i : ’ 
: a» 
tS. 
P* <3 
- = + 
r =] 
¥ <4 
4 
« 
ad i 
as ; 2 tery a 
in eee 
O< 
* 
* 
? 
- + \e >» ‘| 
*, 
+ 
. 
. 
<2) > 
. > 
‘ 
Sac. 
: b Ms 
+e OT) 


aD PUR Vad Bie * 


eckson . ny age at oT 7, “9 
Ketsu SY ecntdos! hate tt) ‘ene 


<i fies a has : en oe ats G bon ‘ 


$0 . itt, EE) 9qgu ate ~seadtt: 


"(ius .a + , ere iW) arrod9 toon lites! 
o*7 7481 ret) - iS & « @) aitah lous (os Lo 7 


r ef wm@liotietea task teeth 
olittieos s dows ,tedel FA ai : 


\ed@iee vileuton doidw Wage 


pre. oPG) «fit acereeR _ eet. ale) oem 


, (eee, ay wafontnwore Lae &, (aed 


4 


Mitoy nie i lege svei- LL ae 

r) , eee, i avelttodl phi 
} be 

tier dye? “ mo Gr S:) Th, 

oat? on 228 


Sike ‘Su bE} 1 orlls eee 


é. ‘tiatieteati org Das 


wel): syteaioee ae 
op sniwehe stemea tt 


*. ans 6 Fe , Saws (5h ao bpryeet 


bas “2 solfer ties) tie Spot é uw 
‘obA oa) of Dono wil fee ohiemses) 


— veeee Sel re, 


—-, Gus ie @. at a an 


45 
from the most generalized pretentoria to those very delicate membrant 
ous ones represented by such forms as Heterocerus (Fig.4¢/9), Endomy- 
chus (Fig.#v),.Pseudocistela (Fig.#, Hyporphagus (Fig.gsv), the ma- 
jority of the Cerambycoides, and the mhynchophore. A very prevalent 


type,possessing a distinctly chitinized cephalic end and a membran- 


ous caudal portion sharply separated from the former, is represented 
in Epicauta (Fig.#sJ, Pytho (Fig.%f, Chalcophore (Fig.wsT, Alobates 
(Fig.#), Bostrichus (Fig.gry), Diabrotica (Fig.#7s), Dendroctonus 
|(Pig.gq), and most of the Scarabseoidea. Rudimentary pretentoria 
are found in a number of genera, as Calopteron (Fig.3%3), Tenebroides 
(Fig.¢z7), Phalacrus (Fig.%z), and perhaps all of the Rhynchophore, 
except Dendroctonus. The pretentoria of the Khynchophora are for the 
most part very delicate and fragile. The difficulty of making a dis 
section showing the pretentoria intact is intensified by the close 
packing with, the snout. of the greatly developed tendons of the mouth 
parts and the strongly chitinized pharynx. In no instance, except 
in Dendroctonus (Fisam) was a pretentorium preserved intact. In 
Scolytus (Fig.go) the pretentorium is evidently rudimentary, but in 
mone of the other genera is there a clear indication of such being 
the ease, owing to the presence of frayed ends, suggesting that not 
all of the pretentorium has been seen. The presence in the Rhyncho- 
jphora (Pigs. 77% 4o9 ) of a suture in the right position for an epi- 
eranial suture, an invagination within the suture that suggests the 


|pretentorina, and a projection arising from the invagination, all 


\furnish evidence that this projection is probably the cephalic por- 


tion of the pretentorium. The pretentorium of Passalus is extraordi 
narily developed. The cephalic part extends as a slender bar dorso-| 


caudead, nesrly half way to the occipital foramen, then bends sudden- 


ist 


5 
at 
J 
>i © 
a 
ih 
‘ 
os 
po 
s ee 
& 
iz 
=f 
rp 
‘ 


~ 4s, if 


eno tt 


x - oe 
oei@) sun s00gb74R sa tmidr ¢onea wde 


oy vier saddéd ot cidetastete Sarai 


4 .- —, 
Mt) aco edie’, tee. ale) Wega 


v 10Soneca 6o7. OAs | 
ne Soe otfedgeo  borind + felt _ Loa , 
,seas0 > adr-horl se regen. | eel 
e924) stoddoplaidc ete ht) one a8 
=“ sifouddid, , soles) eam 
3 .S6DLG8R fwesne Ang to 
me. Le gt ey , teen fo aed 
gti sh, iN e 
: Bryce ‘Io. ai 107 ged ae 8% 


iS qi get ‘oe, 2a 
Joeatnt. as rics medeaeee 


oh ft reern” eat te te 


o ai la 


Feni on nl ,eaurada t ‘auton issovte 


be Sed 
Tan 
& 


PS9AvVE & ¢ vaxt to sonvesay ond tj 


if i iy aost- ertelse ac ieouhorg, er’ * | 


mot otseteyq: a Raw is ae 
Ifvebive el au: Loot teed aig eae 
raefo #& arent? oh oi eey 16 


tees reed aad Fou 


idisoy Jticts adh aL sediwe etal gee 


ay eatatua eg?d wine tw pels spi geva loge; 


ont eldadotqa PL aetiesiomr ajdt tadt 


eeaet 20 cur ligrader ete sat seen 
cette @ BP 2enes ce Teeu » LLadgae aby + 
lie? , meowregs let iglgoe edd of Yam 


ee ty . 
ol a 
i 7 7 

usa *° 
au q 


—— — il ae 


46 
ly and extends'as a huge straight arm toward the corpotentorium. The] 
tremendous development of the caudal psrt is due to the need of a 
strong support for the dorsal surface, which bears a prominent horn 
used in fighting. 


Metatentorium.-- The form and size of the metatentorium is quite¢ 


decidedly indicated by the position of the metatentorinese. Those 

genera possessing Pecmitawe metatentorinae are very apt to possess 
the primitive type of metatentorium, one that is short and simple, 
as Limulodes (fig.3#9), Sphaerius (Fig.37), Anchicera (Fig.¢3y), Philo] 


thermus (Fig.#7), Melanophthalma (Fig.gJ9), and Sphindus (Fig.#¢). 


It is significant to note that all of these genera are very small in 


size. The cephalic migration of the metatentorineae is due to a sim-| 


ilar movement of the metatentoria. In those genera in which the met 
atentorinae have migreted from their primitive position near the oc-| 
cipital foramen, the metatentoria are found more or less deeply in- 


vaginated along the gular sutures, the sutures being the products of| 


|\these invaginations. In most genera the metatentoria advance but 


little or not at all ferther cephalad than the metatentorinae, but 


jthere are some exceptions, in which the metatentoria taper down erad} 


ually, as in Helichus (Fig.w7), Cyphon (Pig.vzz), Lyctus (Fig, an 


jall of the Scarabaecoidea, except Pseudolucenus (Fic.wy. In those 


jeenera in which the lar sutures are confluent on the meson. the ev 
a Ls = 


jla itself is simply invaginated, becoming a part of the metatentor- 
jium. The same type of development has taken place in Necrophorus 
(Fig.3saz), Scaphidium (Fig.3v), Hister (Fig.s7z), Phengodes (Fig.395), 


|Chauliognathus (Fig.3%), and Georyssus (Fig.vzo), as in the Rhyncho- 


jphora. In Necrophorus, Phengodes, Chauliognathus, dupsalis (Fig.¢sz) J 


and Thecesternus (Fig.##7), the line 6f fusion of the invaginations 


Vv | “ee Maley) _ FO a A bait Lee. 


let. tvo°Aerognda odb Bitagoe ave Teeletre 
Hy 7, aceite lob ove) eq Cebu elf Ore 


bo ee be | 
whic = exdad Gitte -soeTrue Leatee “onht 


‘ 7 tigubrovnsbetan add’ to exta oe arin on? aay 
| maitosed)at eu Sat to noitieeg Sim uo Bee 
: foley ae We i: bibl feck Hoy ae Sec avit nedee. | vi 
mor, 4 0 ‘stone oage 
bridl’ otal ateolnema ioe GN invade’. \ (ee 
1, Been tJ Waal am rahe one Tenh - 


; ; A Sal 
‘eed. tL ' es] 22 » toi elon oe +h 
p} : : iy 


Ste. ih Babi ‘afer, ot Qo dell ssl 
le ; : 7 71 ; Ape 
rT Cab ie baa, odd “2 

¥ ; watt svitintre-acene poe saa 
- widgsea) aif ' a. 2 .omelnton edi 
tn | weemhea te } tiod aerscut ue t ,esvoltus “teigg est 
18% 4808 nt: Y enor 

‘eigoo Tedrreh) ote 


fw of -, onesie geure 


a as 


(Wm DEb aanto eee 


i7Vooxks ,eabig 


———— 


oirice taleqg Ode 
hotanigernt &iqobe 


fy noted s inamgofavel tar naget » 


—— 
AS GS ne a 
. 
~ 


reed. Pei ee SGA + es by). tara F asint%) mitodd 
r) ee VIosy ine , eng 


im. nM) i tease arty ensel [rate ,oodouh afi. A 


LE ee A es 
os 


et 


UJ 


sgesltasineyul @42 fo nates oe onl: ate Pa 


47 
of the two sides has disappesred. In Chauliognathus the invagination 
is greatly reduced, and in Phengodes nothing remains but a mere line 
These two latter genera seem to show a greater specialization of the 
gular region than any other genera studied. A correspondingly deep- 
er invagination of the ring-like plete surrounding the inside peri- 
phery of the occipital foramen has occurred with that of the gula. 
The whole phenomenon appears to be due to an especially strong ceph-| 
alic pull on the metatentoria. This can be readily understood in 
the case of the Rhynchophora, in which the elongation of the snout 
would encourage this result. A second force may play a part here, 
that of the narrowing of the snout, which might assist in the enfold 
ment of the eula. 

A large number of genera, scattered throughout the series of 
families, possess prominent projections along the mesel margins of 
the metatentoria, caudad of the corpotentorium, as those of Cybister 
(Fig.377), Reerebis (Fig.397), Glischrochilus (Fig.¢z7), Phyconomus 
| (Fie.#jo), and Boros (Fig.vs). Some of these projections have dis- 
|tinet tendons attached to them, as in Scaphidium (Fig.3%), Parandra | 
(Fig.%7), Derobrachus (Fig.#%), Donacia (Fig.¥#7), and Criocerus (Fig 
473). Prominent lateral projections are present in Lyctus (Fis.53}, 
Osmoderma (Fig.#z), Parandrs, Glycobius (Fig.#7), whynchites (Fig. 
944), Epicaerus (Fig.g#v), and Thecesternus (Fig.g9}. Passalus (Fig. 
$66) is peculiar in the possession of a large well-chitinized second 
ary bridge arising from the mesal margin of the metatentoria. 

Corpotentorium.-- Owing to the simplicity of form of a struc- 
ture like the corpotentorium, but little change is indicated in it 


through most of the genera. ‘the hypothetical type shows this struc 


ture to be a rather narrow simple band (Fig.37#). Such is fairly 


d2tantoavh) #8) sadeprneiingd? of 5h oea0 ms 
Peni ca 2 Jue SB lager wptitton eshognedt at beets 
rm | : : 

bei? to noitorblelssqa> Tereets 2 veoh ot Gaee are 


DAL a= 
- rihneoqesties A&A wheliuts axveties Weai aime 


Hilxue oft So Fads d¢iy ierrstoo san .cematey ee 
i ne | 4) Ch Ve Tiara i 5 s ac OO ty saya: MX 
ait leotre repay vill : Ia? = aivgtaatarene 
troph 8 | resnols Sid ! ,- ody ota gaie am 
Sond ay 
. an’ i ? . £ Pa? oS j ars 7 st igeas alidt: 
Rae 


ioe 
- 
* [—" 


4 “) 
= 5 ; i an ye 

«frog abfetti it agthanoriue of “Ilo ost Rabie ieee 

| 

| 

; 

ah fo tne a: il : isin icscidw ,t#ete si? 7e Bape 


ee 
furan. Ls gopse a! ite a u scotmnny | 


to iiek -« o hetahity se areads Teste 
. 
! bey cy E op Pot mag ins and to Lebsee , 
| | eomor of |, tah. ed id sa lr | (encoun 
f <51h <tavt anedtoclotg sued? torean’ atom eemnn 
what. 4 melita Gc AF oe) eyekié Cee rere 
shore ee AGM | etoenod ‘Rat Bigoe 
| pe SEY ettoyl at ta (@ ano kiest ots Lasetas im 
| ett. eetddonedh  . kw; iidooyte «brea 
} > seeds’ 0 &yesd uated asoaitt hn tg Se 
| peetat? his- iw ental be Ctugeacoqd aise 
. 
( oe fod ae: “ii » nlever Leeun ent (ese atictee” 
~osen. &. Yo mot ts iekliends- edt. of gabeired 2 


: feoltat et eqeeds (La ht pod eldest 
oe 


oni a i : = 
petit? pid? uvede eqyuée leoiteimaqtd én 270.0 gD 
: re, if 


“a 


yiziat of dows =. Gs) fond elaeie, eee 
onenmaaaien ; es 


48 
Se batass 044 of the vast majority of forms. The corpotentorium tai 
sometimes very broad, as in Ptinus (Fig.¢52, Bostrichus (Fig.¢sy), 
Parandra (Fiz.%7), Derabrachus (Fig.%y), Glycobius (Fig.#7), and Eup 
salis (Fig.7g). In contrast sre many that sare quite slender and | 
arched, as Dineutes (Fig.379), Stenus (Fig.3%), Hister (Fig.32), Geo-| 
ryssus (Pig.#zo), and Mycetophagus (Fig.¢37). The form of these latte 
has evidently resulted from the narrowing of the space between the 
netatentoria. An exceptionally large number of genera have only a 
lrudimentary corpotentorium, or none at all, as in all of the Lampy- 
roidea except Trichides (Fig.37%) and Necrobia (Fig.3%7), Hpicauta (Fig 
$o§), Wacrosiagon (Fig.#og), all of the Giidterpldes, Eecphouns (Fig 
Rhysodes (Fig.#z), Phalacrus (Fig.#a), Hippodamia (Fig.gg3), and Ar- 
thromacra (Fig.¢%). The reasons for the loss of the corpotentorium 
are not always evident, though in most cases , either the arms of thé 
tentoria have expanded and approximately met on the meson, as in Rhy 
sodes and Plesiocis (Fig.#7), or the arms are directed towards the 
Imeson and meet there, as in Connophron (Fig.3s3) and all of the Elat-| 
eroidea, or the pharynx rests snugly between the tentorial arms, as 
is so perfectly found in Chauliognathus, where the pharynx is wedged | 
so tightly between them that the whole seem like one piece, all of 
which conditions supply firmness to the tentorial arms and obviate 
|jthe necessity for a corpotentorium. Of the Rhynchophors, Eurymycter || 
\(Piz.4s9), Epicserus (Fig.9#), and Lixus (Fig.¢%s) possess no corpoten 


|torium. Scolytus (Fig.##) possesses a very rudimentary one. The 


|disappesrance of the corpotentorium can best be explained in the cas¢ 
jof the Rhynchophora by the fusion of the metatentoria into one solid] 
plate that needs no added support. The corpotentorium of most of thé¢ 


Cerambycoidea is a delicate membranous structure. A cdmmom modifi- 


- 


4 5 
‘ i 
hc 

, +? 


uiitiethnotaqive oof) Aeaigt to Yaeotem oats 
te, re ). Sst tres. Pe ok Guees ft) eee a we 


RE. 2a) rewil (wet. cit) aucetS eee 


ty» Ue Sev. yee ‘ . 
~ it han Y ie % 4 


ne (eal) sa boomgee ) meet) oti “ 
i c y 


bose. tehhre le ar isi TR Pe “rweay ate taetéaon ¥ 


Eon. O fd . * er i INRSA™O ? eoyls a a 
; . 


IDOE i NAWOLLAn ems mo =r font fare = 


[na ovubtlmeches Yo rede endear yf! snobt Gea tbe 
ner So. a dota ¢ e 

tse! Lgo toe! 5.90 sondtod 

hr ; foreterla eae le: Tia restyled F 


fete b ai ougih , git. 2tt) ase Ree 


oy ; noases ont 
= a : “4 
m ; . FSO wi eoarostt , Ba sares Ne. 


rane ree bobaasene 
wot  oyeo oe 40 (eueate) aoe 
r‘Te G10) neotsoamnire nr: voted 
ssvieu QE Bl eo eee 

oifvents mi Ripot ¢t 30 


soe] vate i slortw sig fang, wei ies 


Ty Se 
x 


Ewe wm 3 Colitodbet sat of beanie ‘Lage Hees 

a paongan” 4 tO 6.mvivgtaslogt0g a gee % ; 

Ge hee. oll) soxit ide , AWe. alee eurone oA 

ede gestieantios «sey n seneemson hii a) 

iuad asd owliofastoatig. Sa a 

ond ofnk elpotostatec ant %6 Ao leur aie so 
¢ ovlwotsetoyreo séf) Gage wea 

, 


N i iNT 4 4} ori ‘ ra SC AE ay ’ ba . , 


“ghee ew eee 


49 
eation of the corpotentorium is the mesal projections on its cephal- 
ic border, as in Tachinus (Fig.3%7), Cephaloon (Fig.7o/), Tomoxia (Fig 
43), Phenolia (Fig. 4s), and Philothermus (Fizg.¢3s). 

Leminatentorium.-- A structure of the tentorium that shows per- 
haps a greater variability in form than any other is the laminaten- 
torium, which appears in a great array of shapes and sizes, from the 
forming of a perfect and broad bridge to total disappearance. When 
the two sides of the laminatentorium meet on the meson, the mesal 
margins of the laminatentorium very frequently are bent ventrad, pro] 
ducing a projection. Examples of this development are seen in Calo- 
soma (Fig.373), Cybister (Fig.377), Necrophorus (Fig.3s2), Aleochara 
(Pizg.3%3), Heterocerus (Fig.wy7), Dermestes (Fig.#23), and Endomychus 
(Fig.7yv). Genera that show the leminatentorium as hardly more than 
touching on the meson are Peltodytes (Fig.3%), Scaphidium (Fiz.3y), 


Pyrochroa (Fig.#0j), Notoxus (Fig.#/e), Psephenus (Fiz.¢%), Cucujus 


(Fig.7y), Penthe (Fig.#s7), and most of the Scarabaeidae. The reduc- 
tion of the laminatentorium takes place so gradually that it is hard 
to tell when it has completely disappeared. There seems to be no in} 
dication of it in Limulodes (Fig.77), Collops (Fig.3y7), Pseudociste-| 
la (Fig.gss)}, Hyporphagus (Fig.vs0), Sphindus (Fig.vse), and others. 
lIn Tomoxia (Fig.#3), Pytho (Fig.¢07), Alaus (Fig.#/z), Tharops (Fic.9 
Throscus (Fic.¢¢), and others there is no laminatentorium. The pre- 
tentorial arms curve towards the meson and serve a similar purpose. 
The laminatentorium of Tachinus and Epicsuta (Fig.go9} is in the form 
of a lobe. That of Psephenus (Fig.4%), Dichelonyx (Fig.75¥), and Pel 
idnota (Fig.70) possesses two long sharp cephalic projections. 


Supratentorium.-- Arising from the dorsal surface of each pre- 


tentorium cephalad of the laminatentorium and usually extending 


2 


wa *L 


wll 


mat 


ane. 5/8) guabdogad ween be ovens Oey 


sores , Ge.a2t) soets , hebiett) otgee 
mesrotoetaning! om ah voxedt or0edto fie 4 ae 


at 


Weowtt) zyooledeltd (aes te) ceoert eg 
.aeoli sealer fedqan qaede snot owt a4 
done to coghaue Laethd om? agit ee . 


a6 aneiteastdta Glesa attied mn 


off uf weatfo (ne 2sde eee yeriid 


sta ine seus 2O Yetta eee 
w“agoreei— toto: of es iietibseord: Bieter 


ih bas: MOS erice © saTy bor het. ‘yale 


af{tmin a e7tee *ee nodem. eff. shaewos evtyo 


4. twe.2ll) etateotc hos aontdoat 26 aartotae 


Je cit) sueiegoed RR eaeey a 
» fee, cee) Shine: Lette yous 


d+ mabsotaey ea¢ te oat oaale 5 ab 


gut : 1G -T QO ovirotnetenings edz. - . 
Me sae “lteenposl. (ievees totaer am 
tommetfLeves aide “*oelgmease £0 

i 
aA S ecle  a vwEoruotgar detund@) Tee 
tse. 2th) sotuaaed oe iy 


nubtotns?éer inal ett voite a 


la se ss To Goon 2u8 ery 5 4 

% | 4-aagiq secat wes Protea ; 
yreqgqeal b 4 od eID ps part 

hat. 228) agotled | iyi. ai) aebe tae ma 


ta 6, awa IE) suheisgs , ee sit) 2 


50 


towards the dorsal surface of the head is a projection of variable 


Its generalized condition would show a ra 


form, the supratentorium. 


ther distinctly chitinized structure, with an expanded ventral end 


against the ental sur 


that gradually narrows, then expands flat-like 


face of the head-wall. This kind of structure is found in a very 


large number of geners, as all of the Adephaga except Peltodytes (Fig. 


37%), the Hydrophilidae, Leptinus (Fig.3ss), all of the Steaphylinoidea 


except Aleochara (Fig.333) and Hister (Fig.s7z), Pytho (Fic.407), Noso- 


dendron (Fig.vzs), Languria (Fig.#3z2), Megalodacne (Fig.7s3), Derobra- 


chus (Fig.¢s), and others. The supratentorium is a surprisingly per 


sistent structure considering the large number of genera that possess 


it in a more or less rudimentary state. The broken ends of the ten-| 


toria of Hupsalis (Fig.¢s2, Lixus (Fig.+s), and Sphenophorus (Fiz.+s9 


_|jare expanded, which expansion may include a part of the supratentor- 


ia. This seems reasonable to believe when a rudimentary one is evi- 


dently present in khynchites (Fig.##) and Attelabus (Fig.v7s). Sphen- 


ophorus is peculiar in having the lateral margins of the tentoria 


fused to the oculata. The head-capsule of this genus is suddenly 


lconstricted at this point, which probably placed the tentorium and 


head-wall in contact, a fusion finally resulting. 


The supratentor- 


is of Cucujus (Fig.¢3/) and Passalus are stout structures, but ex- 


|jtremely short. In the former, this condition is due to the flatness 


of the head, in the latter, to the unusually close proximity of a 


jpart of the pretentoria to the dorsal surface. 


V. SOM PHYLOGENETIC CONSIDERATIONS 


With the hypothetical type of coléopterous head in mind, it is 


possible to note the lines of development that have taken place with 


in the various groups. Structures have developed away from the prim 


iF lees Ser oem aie Sebcetties.: 
ofdeinay To Roltospesg @ Ob baed ed) 26 ao Cea 
=a vole bivow oor higep hegila tereg a2 i mt 


<toev. bebpages. ah titi ,erotouitacbeae 
-~ : , an ’ bate haan an iif al ahusyre sod? 


tie to bets aige= 


>" Patel sox piiovnetaass “wg bast, 
‘We BA RO Pr teadd? # cen 6 sadn savel  6e2ci wer 
~Ner ant De 6 [ LOT ; ,e7 az Cth at nom DON ae we” , 
E : “a 

Kes, i) as none Caer nh) bivet I: a 
isinel arom od ; y @ obifoni ¥y cui tolenadnns 
Hine i + S&L ‘on pe a ‘gts aa@ weoekad og cs 
Pes enae seen tl) ascdalorrs RS \e. ule) cor tena 
by nebes > ni a. Savetog ond galyary ‘ab 
. gineibstg «i Loy rai ) olWwaqac-peen aise: 
bre ovitoevass f cata ufdedowd Geteiw : ao ‘enat 


-—sofgeTetaus 9:7! wih? foget gilsaé? ce test & ., Fon? ¥ 
J ee 


= tO) io. ., Sa vie fuved@ B12 SA SG. bom ie 
: 2 psrh } (OAT L Daa sis’, Tons elit awe 
O et imixoze japte «i Leweuns mf oF ,corees ont 4 
ooatine Lento ems a J strenneny 
LOTS ASACTAKOG OTe RuRe ES Oe, M 
oE ti Soto al Saod eootetapsita B6 egy | leek 


qd lw goely cudad ovat tadt. + neago Level to: aemks 
ely cy ail not} ma wa pomp Leven apes soTintouspe - 


51 
itive condition with varying degrees of rapidity, and sometimes in 
different directions. It is difficult from this study to decide on 
the amount of importance to attach to the change that may take place 
in a particular structure, and to average the degree of primitive- 
ness aS a whole of the development of the structures characteristic 
of a group, and to indicate with precision the place in the primi- 
tive scale of each group. As previously mentioned, owing to the 
wide field covered in this investigation it was impossible to find 
time to study a large representation of genera within each family. 

Such a study should help one considerably in reaching clearer con- 
clusions as to the arrangement of the genera in the primitive scale. 
Using this study as a basis, it is quite possible, however, to dis- 
cuss the relative importance of the structural changes exhibited by | 
the different groups, and to suggest possible inconsistencies in the 
present arrangement of certain genera. 

Adephaga.-'The characteristic possession of the occipital suture 
jof complete epicranial arms, of a generalized position of the anten-] 
|nariae and of the pretentorinae, of a generalized form of labrum. and 
of tentorium, would force such families as the Cicindelidae, Carsbi-| 
dae, Amphizoidae and Omophronidae into a primitive place in the sie 
jlogenetic seale, in spite of their specialized ventral surface. No 


other large group showS as many primitive characters. The genera of 


|the above-mentioned families show a great similarity in all their 


istructures and must be closely related. Of these families Omophron 
possibly possesses the most generalized dorsal surface, though the 
Joccipital suture in this genus is not nearly complete, and does not 


|show on this surface. Omophron can hardly hold an intermediate po- 


sition betweem the Carabidae and the Haliplidae, for the latter 


ets 


fnmihenge foe ee Sieges bo Beetyen: ee 
ofituak of Cbd arte mpi hire Pytus et th) " 
| yan tes Gyenep out ot dog ts of 
viftininge te. sovgeh eat oaateve oF Bae went t iy - 
frafoatan® esigveriTe: en y 56 Prenyo lowell ade 
heb ost ot, aoet 2 soiebors r oy iw, etaotbis 
i> bf wdotwo ,senoti ~€ogo0fvesg as quotes 

it of aidiveouwwt f4n D4 no lvante b sri ni 
fima't dome didtiw asczres to oot atneseiiee one a 
eeiveth ipifess® cid vidsaetianged agg gles 
ne avlttaltew oof o Yeuen. it) Le Perry cr 


Bhi hae ik ints a ee Se ee 
Pe: i 


wd #efivtinze sovoecos. Laralouate ats 3e osned tog 8 
™ ’ enefaicnoons 4] 7 teens ro mre Hts’ X ring + 7 
avaiee a lettea te teen 
B! qaubaey ee | iaadoaog™ oa tretogtaite 
= y To iPhean hes Pieters eRe ere a Tad 
aitere’ sa) Be fsisn @ Te exirosaosiong dvi 
Litured!  papifevulor ed¥ ap tine toot onthe prow j 
fl sae me eoaety ovitiwita Bs etfag sab nos oind Poe on vik 
«90% ed extn Leo t Os boas Slate te ofiqe (ah a 
~onen at? .ouerce risen evid inicaeyalee 62 ewe 
fle nd fap f desta etn 8 ede setituwt be 
ciqomt SeFL Tm ott “0 -Leteged® qtetete ee tom 
{2 soit ,eoatrtie leatoh beet fetenay Gem ‘one aqunonad 
He .jote lie VTSSs. tot ah Sipes ated wh. salerbang 


weliompetat ae £fol¢ heed eae nope casei $i 
Lod) ef aii, To. ae 


52 


latter family, as represented by Peltodytes, shows little similarity] 


with any other family of the Adephaga. The form of the head is dis-] 


tinetly specialized, being globular, with enormous eyes, and the 


mouth-parts and the very small labrum are directed distinctly ven- 


trad. The occipital suture is probably lacking, the epicranial su- 


ture is shorter than in any other genus of the Adephaga. On the ven 
tral surface, the metatentorinae extend only halfway between the oc- 
cipital foramen and the submentum, and the gular sutures extend no 
farther cephalad, whereas in all the other genera of the Adephaga 


the metatentorinae are close to the submentum and conplete gular su- 


|tures are present. Finally, the tentorium of Peltodytes is not so 


well developed, and the supreatentoria are small and do not reach the | 
ental surface of the head wall. In all other genera of the Adephaga 
the tentorium is large and strongly chitinized in all its parts. The 


Dytiscidae and Gyrinidse are very Similar fundamentally, and differ 


from other adephagous families in the position on the ventral aspect 


jof such structures as the antennariae and the pretentorinae, and by | 


a rudimentary or absent occipital suture. The instability of the 
epicranial suture is well illustrated by the case of the two closely) 
related genera, Dytiscus and Cybister, the former possessing com- 
Iplete, distinct arms, and a considerable part of the caudal end of 
|the stem, the latter incomplete and rather indistinct arms and no 
art of the stem. The Dytiscidae and the Gyrinidse sare undoubtedly 


Jmore specialized than the first four families of the Adephaga. 


Hydrophiloidea.-The form and texture of the head of the Hydro- 


| 
jphilidsae is quite similar to that of the Dytiscidae and Gyrinidae. 


|The heads of all these families are broad and short. and strongly chi 
| 
| 


‘tinized. The antennariae and pretentorinse of the Hydrophilidae 


Avtiaol bel: 


Cl IS TT OT = 


a A T 


it 
ad 


¢ : 


a 


sf a 


¢* haw Sl 
y 
t | 
| 
(4 
° + 
ney bh ® b ‘ 
‘ ' 
A 
‘ pa s@#bi 
Ta Boe (See 
* di 1 aur 


choo wou aati q iad a o 


te say net caer’ 200 ae ze 
bat ed "daanie real martin, 
ween i ete, fib 
bnetixe GRs lrotnetedoms 
ft 1 et in a | 
<9 We Ca 3 asexoy 
ad agate ona’ 
¥: at ae an caatt: 
i Poteet aia ere 
eacle fant. La be ont “vit 4 
il Dinnone ine sereti 
be selinda ype ate, osbiabrys 
preted. Shit aed Ommeane eurogag 
-b estranaelRe, ont uae 4 
etait led totese Samed! 
1 va 5 teabn: | et EE baw 9 9 
“iy ,rotmeidye Ose ached git gil 
eidetsh Foace 3 ana aes a 


iiet 4 10 ay wT ee A lqenopad tadt +f 


tiv) ot ga cebloulhedl oF 

‘tfimet toot tetkd odd) all 
i to exudect Bae ereh eats 7 
itya-lt Te fadsunt 24f Dake. 


ine ones? ote se iflaat, 


53 
have migrated ventrad, as in the other two families. The metatentor 
inae, the gula, and the tentorium are almost identical in form and 
condition, also. This superfamily is evidently very closely related 
to the Dytiscidsae and Gyrinidae. 

Silphoidea.-- The genera of the Silphoidea show many inconsis- 
tencies. The dorsal and ventral surfaces and the endoskeleton of no 


two genera are nearly alike. Necrophorus possesses the most general 


lies of the Adephaga. On the other hand, this genus possesses the 
Imost Specialized ventral surface present in this superfamily. The 
|guler sutures are couplete, and for more than half their extent meet 
on the meson. In contrast, there are practically no gular sutures 
in Connophron, a scydmaenid, and Molamba, a corylophid. No greater 
extremes of. this surface can be found within the limits of any other 
superfamily. The tentorium exhibits just as great extremes in devel 
opment. That of Leptinus is quite generalized; that of Necrophorvus 
well developed but specialized, owing to the invagination of the 
jgula; that of Connophron with no corpotentorium nor laminstentorium 
and rudimentary supratentoria; that of Molamba the most rudimentary 
tentorium of any genus included within this study. No clearer illus 
tration of the difficulties of indicating relationship can be shown 
then that exhibited by the families of the Silphoidea,. 
Staphylinoidea.-- The various genera of the Staphylinidae show 


jconsiderable homogeneity. Tachinus departs the most from the charac 


ized dorsal surface, which is very like that of the first four 
|teristic condition, in possessing a much more generalized dorsal os 


|face than the other members of this family. The Staphylinidae poss-} 


jess a tentorium and a ventral surface similar on the whole to that of 
| 
the Adephaga and the Hydrophiloidea. Tachinus, through the added | 


asoticn fet or 


Hie wre 


ivtat yey ot 


i) 
: 
ml 
i 
= 
caw 
— 
, 
Sat 
at 


a 
i 
: 
Pr) 4 
~~ 
m 
~ #! a 
B-Lztss 


oq anbiolizdeedd edt. Rie aad tee cueine 


Depa 


A ebbinek, ‘ow waste ‘ath at a 


ovtéy hea bes 


‘Joa brig tatthor bua 

irtetge ee nneaes vt 

am mone teh esha ¥ 

| i Load Sane: ay ag” da) 

feétral tae ao Werte, Daveige 

Led: Qaits Saou Bhi | abstgnet 4 
aiiin, ¢ 7 Sifae ta Big ostekbe | een 


el ahs fyvroo Ay atgvelom Lae, Akar 
Pn 


f a 


£5 eter 2 Le \ ! Deora ish ad BO: one 
aiemartice 1: a aa Peg arate. satin 
seating p eget er ii op, ot tage GR suntan 
& hlltncbnevnds odes Oelee boa ssatooane’ 


Ho. 


iobrot ned oe et pears 
old PdmeTotisko afm cl eaieie 
obots eit). Reo econ saa 
- of ha oe an ite toed ae wottts 
ecdwete lat we iin 


r 


ieticnee’G aff oo wiansgegehear eft eeg 


ip moe? fa0m anc 2!iaqgsh BRres? > JF : 


f asile ene, eton Oem 6 nrtepapnsog nt 


ietive ei2 Ao ‘oPleds ete ltu® ifaw 


‘off isvort! wie pon ek» + seote meh aaa a 
2 


a ae 


54 
similarity of its dorsal surface, would seem to bear a particularly 
Close relationship to these groups. The Sphaeriidae, as represented} 
by Sphaerius, and the Ptilidae, as represented by Limulodes, possess 
@ primitive condition of the metatentorinae, while Scaphidium and 
Hister possess confluent gular sutures. The Pselaphidae, as repre- 
sented by Pilopius, in form of head and character of the ventral sur 
face and tentorium shows a close relationship to Connophron. The 
irregularities of structure displayed by the families of this group 
are nearly as great as those displayed by the families of the Sil- 
|phoidesa. Both of these superfamilies probably need considerable re- 
|vising. 

Cantharoidea.-- All of the genera of the Cantharoidea possess 
complete epicranial arms and dorsal position of the antennariae, and 
lack a corpotentorium, except the Cleridae and the Corynetidae, rep- 
resented by Trichodes and Necrobia respectively. All possess a com- 
plete gula except the latter genera and Collops, the representative 
Jof the Melyridee. This superfamily can be divided into three sub- 
groups. One group will include the Lycidse and Lampyridsae, charac- 
terized by a globular head, short,broad gula, and absence of the pre 
tentorinae. A second group will include the Phengodidse and the Ca 
tharidae, characterized by a longer and flatter head and longer gula 


|The third group will include the Cleridae and the Corynetidae, poss- 


}essing short incomplete arms, incomplete gular sutures, and a well- 


|}developed corpotentorium. Collops does not seem to fit in very well 
Janywhere, though this genus, through the form of its dorsal surface 
jJand tentorium may possibly lean towards the second group. Cantharis 


}of the second group, is peculiar in possessing s normal gula, where-| 


as in the other forms the gula is rudimentary, though in 911 other 


Pietra cee y ei v ei F her exiyl eat 7 ae 


é ec amo Tat if Hers wtdors iat Mere ey 
§ f j Ma gst - f ty vim . eg fy hae ey 4 % uPeG ate 9 
— aa. au 


nottyorng!. 0% Widens ais aang a ewoila.t 
Se ar ne a es nh ch ab ve be vale bb Sra pureed t 
os RE eh. hs | bd Peay PY. Fon seyekas ss “eee aa | 


Bes! edt hag sphlivopned’ od shategt Sige + aarp i 
Refon qeanor.t - feod xs ; alt Sas trepgol & aes wt 9 
biteoeied odd. one Sebd7eld lott, She stiw 

fi » foe sevivtsce talun wr elaroort + Sent otek 
ice vie? al tit od osese don 2eeal got Loo este af 
ong =7 fantoh ail fe mrot oft topotal jaaeee “ae 


itedédanod  .acowe Dgénen ede ebcewet’ neal adr 


ie Or W ‘ ayy | ft SET LCT 4 Bi ] aaedeod ot ish taong oe 


gitito [hei jab ag ak dan | 4 5at wiry 


55 
respects this genus is like the other members of this group. The 
first andusecond groups sre no doubt-closely related. The latter is| 
probably the more generalized, possessing a dorsal surface resembling 
to a marked extent the dorsal surface of the Carabidae. The gula, 
in so far as it is complete, also resembles the condition of this 


structure in the families of the Adephage, and particularly in Necro 


jeven, than in Necrophorus, being not only invaginated, but reduced 
| to a more or less rudimentary state. The first group probably e- 
volved from the second by the change in the shape of the head, due 
to the enormous development of the compound eyes and the loss of 
much of the gular region, which has probably fused with the cervix. 
Parallel with these changes, the pretentoris shifted their position 
so that they could better support the globular head, and came to as- 
sume the direction ordinarily assumed by the supratentoria of other 
genera. The third group would appear to be more closely related to 
| the Mordelloidea than to this superfamily. The condition of the epi 
cranial suture, the position of the antennariae, pretentorinae, and 
metatentorinae, the degree of development of the gula and tentoriun, 
would all favor this assumption. 

Cupesoides.-- The representative of the Cupesidae studied does 
|not show any particular peculiarities of structure of the head that 
| would entitle it to be placed in a separate superfamily. It can be 
Peery readily included with the Mordelloidea, for approximately the 


| same reasons as the Cleridae and Corynetidae. 


phorus, of the Silphoidea. It has, however, developed much farther 


Mordelloidea.-- The genera of the Mordelloidea show considerable 


|homogeneity. Most of them possess hesds thst are elongate, with a 


generalized dorsal sur 


eaf acon fut? Fo ar atiron recdito echt cxnd 6h 
fei tottwl eal, -sbaviedes Ciheols: ¢ aes ae. evel 
‘Peildmevet goetuen featohie sirfoasseog ben 5: 
ofes sdf .apbtdegead eae. to etetiey fantok 


pind to nett iinoo’ etdivesl dese: opie (¢ 


, tai a gouia Tewoleveh ,t6everor en. OL wy 
bLedpier tod, .feovearineatrt vino ton wa ted i iter | 
art heat aft egéen , } grado Omg yd: Saoce 
L¢ of sig ¥ ifourids. SAG.) Te iremgo 
xi of «at > ia Por be gada 4 * 1a Siw , 26 5 
1 soit og. Tiedt J + ite ag wend ary ibe i 
Bucs qt “Be hie ¢ Sn “reebsaod- fe F | 1oogae sate 


godt’ to wivotinet 215 + og psgateeas ifiteete 

| os psd: fee wieaofo 5461 dé af tpt DbigowW quo 

alae 0/2 Od? Lone > 6c vf inet teqge’ aids fo ‘ 

: ~ - 

bie- eeafrofnas stg wf itsnret nasa. Fe noltiecg ede: 

Pwrltotne?: Diu m ont 3¢ remgotaveb to oot wit 4 ; 

, Role Qo —? 

ooh beibuce eal be oan aid LG ovitet nes esqeanedtes 
oft ied ott De eigicaita *o ee hhieet Pros tees 

é ms 27 ‘y fins ‘thyv® eftegqeqns ao of hepate od ot ial 

hx temtroegad tot ,2ebicissiiol axtt ae tw, 

vs 

cobltongiot tee sabitely odd, 


Wdrieitans» sofe sebioflotiel Gat =a s.8n@g; eBt, * 


6 dijw ,efepacts ow tad abead: (gated spedde ike 


56 
on the same general level with the rest of the dorsal surface. The 
epicranial suture, however, shows considerable etenittey) sometime} 
showing complete arms and stem, as in Epicauta, while in others the 
epicranial suture may be extremely reduced, as in Macrosiagon. If 
these two genera are at all related, the condition of the epicranial 
suture can hardly throw any light on the degree of relationship. On 
the other hand, the metatentorinae are stable in position, and are 


located almost uniformly about half-way between the occipital fora- 


tH 
if 


men and the submentum. Another characteristic feature is the fact | 
that most of the genera possess heads that are distinctly constrict- 
jed at their caudal ends to form a neck. The Oedemeridae, the Cepha- 
loidae, the Pyrochroidae, the Pedilidse, the Anthicidae,. and possib 
ly the Cupesidae and Meloidae, seem particularly closely related, 
through the possession of similarly shaped heads, whose caudal ends 
are distinctly constricted, of a generalized dorsal surface, of com- 
pound eyes that are usually more or less emarginated, of a ventral 
|}surface whose structures are practically in a similar condition, and 
jof a tentorium that is alike throughout, with the exception of the 
Meloidae, in which the corpotentorium is absent. Though the dorsal 
surface of the Mordelloides resembles to a considerable extent that 
of such families as the Carabidae and Cantharidae, yet this surface 
|} differs so widely in closely related groups that we can hardly place 
Wess much importance on the resemblance here. What seems to be far 

| more important is the totally different condition of the ventral sur 
|faces in the Mordelloidea and the Carabidae, for instance. In the 

| former the gular sutures and the metatentorinae extend uniformly 


|}half way between the occipital foramen and the submentum; in the Cark 


abidae the gulsr sutures extend the whole distance between these 


a ie one he feateb ut fo. ae wre nd es 
pind aoe. ome cited abt etdateh thao smal. 
ett ansdto ot af fiw shade nls rll | 
. sndaste omad af ee emanate. 5 fomegtine oe bs 
rathavotge ef? te poletiwes of) ,codake iene ale 
aC. .qidenottate bo verges is m6 ‘elk Rem pmol. 
| e Risa bolt hou fi eldats 3% sotrebaedad sar eat 7 
| wate® Lara iPioo sii aetod ewe Lod dedi vliretane : 
'‘toat gute et atirts liek sated redtoad. . muPevest 
Bite betaine Sahel fest Sheed Ssagapeg atenapg 2 
nye bro ot ehne:, 
now bam) ,eabioind piir ,ash ht hoe petit eabie nh } 
beer, frei! we oebroLem aint 
: ores a 
eeode , Glas C ie 3 | poe 29 nO Diwe 3 
“a> %o...aee ines ret Lcahegeyie to Stet 
meh fee ra. ata cRine gal, do ALOM eh heumas otee teak 
ot? Lhe Af hake ot wWilewtlieete ore meted olent am 
te aekiqesre, ett colo deromsuoude ew ite at das? 1 


im t -j2ahde @! cebvediet ogteg ade Cote 


ta ‘Ars oltanshlenodo a of 39 dmen OF ooh 2oL.Loneoth 
¢ “pieinitt S6r . ; sifnad) Bae sabidezeo edd ‘Be 
\@ . ‘ ute pend: TwOTE hetalen yleeafo a2 eg 
; 4 namee ire] ered Soe fdmet et ony ac. pomett r 
y oft to soli ibade teeteti is yitntet one mt ¢ 
» en eourlegct vot ,enhtdaetad od fee oniarinee 
tian Saetts eanixotaeteten sft) Jee eormdue 

iL jouw tenc@e OS fos nameto!s Lattysona en i 


eset cnentved epgeteii afore age bases 


YS 


ah 7) 


57 


parts,and the metatentorinae nearly to the submentum. The Mordell-| 


oides and such groups as the Adephaga and Hydrophiloidea do not ap- ; 


pear to be nearly related. 


Elateroidea.-- The members of this superfamily are homogeneous 


with the exception of the Buprestidae. The pretentorinae and the an 
tennariae are located on the dorsal surface, and the epicranial arms 
extend no farther caudad than the pretentorinae. On the ventral as- 
lpect, the metatentorinae extend but little cephalad of the occipital 
foramen, and the gular sutures no farther. The corpotentorium is 
lacking in all. In the Buprestidae, on the other hand, the preten- 
torinae and antennariae have migrated distinetly ventrad, and are no 
visible at all from the dorsal surface. The gular sutures extend 
the whole distance between the occipital foramen and the submentun, 
and there is a distinct corpotentorium. The Buprestidae are hardly 
closely related to the rest of the Elateroidea, 

Dryopoidea.-- With the exception of the Georyssidee, the member 
of the Dryopoidea appear to be fairly homogeneous. The chief charac 


teristics are very prominent widely separated metatentorinae, locat- 


1 ERI ES 


ed very near the occipital foramen, and a very broad submentum. The] 


ventral surface of the Georyssidae, as represented by Georyssus, is 
jtotally different. In this family the gular sutures extend nearly 

to the submentum, and are confluent through practically their entire 
jextent. The metatentorinae cannot be definitely located. Georyssus 
|jwould appear to be related to Hister, both through the condition of 


|the gula, of the tentorium, and of the dorsal surface. The general 


| 


Jappearance of the ventral surface in the Dryopoidea is much like that 


jof the Hlateroidea, and in other characteristics they are not unlike] 


These two superfamilies appear to be related. 


wt 


fexiiny ton ote yer ach ottetsennts taido, Rt ge 


pieod ef. to De lenges .S2. J i a wa smote. 


He 


iT 


Abt OM oa? hie eae ot feet i Oy 
for of a ead) Fok ililg th ‘Paw & dash a re 


rava omert oni vl inet: <egue aiit ro one 
lt fue ona trot oat oxé oso Joh rasntall oa 


fed oe et rg S auth She. , BOF rn tcx > Spee ral ‘eit Le ™ 
py Pay acl om 4a leesneT akg add ian bebuge: 3 


(irotast ogiuwe' ott told ret on nonitwe <olay. 
wy add abaas “ta! BY) ROU chicsetqat ao oat a 
bea .hetrney piteatias atime oved eats 
arte ee rsc POLin wail = rise teerob vat 
pentite eff fra ocsohevol fate Satan exit: 300 


bitserawe Sil saerts cused 


+ Hebi ors ya tat eat Xo 1008, sai 
— ee of oct. te apetgeors ail? arm os 
tio 8 a £euensxn omen cidor ed OF, roe 
pol .senttotustatom bei ertegse (febie rym snot i 
mitiendus beord ¥; o7 # big ,;Remetot Sat tgteoo,, 
‘ayToAO ¥ , <6 Qn ind Lng esom® eat reg 
im foolx: eyotis Sh Coe enc jiinet pert: ae 
: t*led!’ vilooltos 14, cpeoTay teanftace ote bas Ph. a 
artoch§ vhetenol | codentvad ed Jongao sanltorn tad o4 
roitl brat wl 7 ssiota? d?od ,dvers Lh, oF batalet ndiet a 
£ 


as od) .eottane Leerof ont te, baa estrotaet Wit to 


dif nonm ea! aab ing’ raxG ad nds ooektee aT Ld A 
Sa 


botalas ad of bina sag 
= ~~ 


— a —_ a - en = a ee 
e fii) eae —— mo re 


58 


Dascilloidea.-- The two representatives of two ot the families 


of the Dascilloidea studied are 


Similar in structure. There are no 


outstanding features that would 


warrant placing them in a separate 


superfamily. They are negative enough in their characteristics to 


fit in, perhaps, in a number of other superfamilies, such as the Mor 


delloidea, Cucujoidea, and Tenebrionoides. 


Byrrhoidea.-- The representatives of the Dermestidae and the 


Byrrhidae studied, Dermestes and Byrrhus, do not seem to differ de- 


ecidedly from one another, but the Nosodendronidae, represented by No 
sodendron, may fit in better in some other group, such as the Dryo- 
poidea, 


They resemble the members of this latter group in the posi- 


tion of the metatentorinae and in the form of the submentum. In No- 


sodendron, the metatentorinae are almost adjacent to the occipital 


foramen, with a membranous sarea included between them. That part of 


the metatentorium surrounding the periphery of the occipital foramen 


projects deep into the head. The supratentoria are well developed. 
(None of these characteristics are possessed by the other two genera 
studied. 


. Rnysodoidea.-- The representative of this family studied, Rhy- 


sodes, shows distinct peculiarities in the structure of its head, 


such as an extremely thick chitinous head-wall, a fibrous condition 


jot the cervix, an errow-shaped head, enormously developed submentum, | 


jand very small mouth-parts. Its relationships are not at all clesr. 


Cucujoides.-- The members of the Cucujoidea show on the whole af] 


short broad head-capsule, a distinct ventral migration of the anten- 


jnariae and pretentorinae, the degeneration of the epicranial arms, 


Jand a slight migration of the metatentorinae from the occipital for-|} 


amen. There is some variation in the dorsal aspect. 


The genera 


u 


kh : i " of wc i - ¥ we: Py Ti 
A | 7 7 } 
Poa a a 4 
Kies . 


P ralliok® ead 26 ont te sevitetous tq ont 


y , is - 7 ~ 2? De Uy ¥ 
Ree emit te | fs or: 


ad 


. 


ae | 


6c ot e1ontT ‘letatnamhe OF myliote are be bbs | 
s3709 e4ht mod seRny ¢aester BLoow sade, Q 
treltatobyredo wheds AL durord sv itegen ate ¥ | 


= % ae 
col’ sant ee Nore ,eadlinetredgs todto fo Tegeo ae 


Tae a” pnp na liom 


S ter esebitesreretd edt ‘ty joy it ers ASI¢tSes PY, eae 
. -. =e" fed 7 ee 

rotrinp oF meses FON Gh ,t igtCe Ona y eve sarred ele 

of. vt befabesitdysy v £ trom Gae Oar tad , TOA Cae 


/ e f 

+ 1! satted of ttt? 
| path 
eisog o! if “dot tettef elit to etedwam ge wie 


a i . sor Mmemnoie a5, iw i AF conixotpenad 


> 


ole ets oon tetas 
mig g o4 if omant peewted hebufon! rors svocatiiog 

0 lq ited ed? seiisqwornse vie 
sh is 
sy F I y: eTdUea Uod ,fsen and of eg “ 


is 


_ = r . ¥4, a 
aeaoke STA toitalrefee reds 


i 
- ts vitatasestqes eng 
, i ab gage ts Lies epates 
hy i ret v- Dead @ ton te Phe ‘tots Ri ome * 
: sfog eqgolevel) ylavomtene ,baed begastamaure ne : 


Ths! taf » polugnottefesx agg at tog-dtwom OE 


oe 
ve 


P Y no roid aeticloor? add to Saggy SAL seen 
5 4 ' toliatminm lArtaev tontteil Se oLimnyncsiged oa 
— 5 Ineteiqge off to nold@a éqegeb eae santvodwsstes 
Pye * f[etiqione éa¢ govt eastltor asset sm aig ba) Saierordiy 
. Zieves act 1) Baru Lgeioh edt sb aokteks ; 


vm rat - hay | 


59 


belonging to the Mycetophasidae, the Mycetaeidae, the Melanophthal- 


midse, the Endomychidse, and perhaps those of a few other families, 
possess dorssl surfaces that strongly resemble the dorsal surface of 
the Mordelloidea. In Derodontus and Philothermus, a colydiid, the 


antennariae and the pretentorinae are botn on the dorsal aspect, lo- 


cated considerably caudad. 


The ventral aspect ot all the members ot 


the group, however, is strikingly similar. The tentorium is fairly 


uniform throughout the group except in Tenebroides end Phalacrus, in 


|which the pretentorium has distinctly degenerated, due probably to 


the heavy chitinization of the hesd wall. In the shape of the head 


6nd the condition of the metatentorinae and the gular sutures, the 
Cucujoidea show a Similarity to the Mordelloidea, and to the super- 
family Tenebrionoidea. 
Tenebrionoidea.-- There are no particular characteristics ot thé 
head-capsule that would separete this superfamily from tne typical 


Mordelloidea. For instance, the dorsal and ventral surfaces and the 


endoskeleton of Arthromacra, Pseudocistela, and Penthe, are almost 


exactly like these parts of such genera ot the Mordelloides as Epi- 


ecauta, Cephaloon, and Macratria. The corpotentorium of Arthromacra 

and Epicauta is, however, lacking. 
Bostrichoidea.--The structure of the various parts of the head- 
jecapsule of the representatives of the Bostrichoidea veries consider- 


jJably. The epicranial arms are complete in Bostrichus, Lyctus, Plesi 


ocis, and Sphindus. With the exception of Sphindus these families 


Show a distinct ventrel migration of the antennarisae and pretentor- 


jinae. In contrast, Ptinus and Sitodrepa do not possess complete 


jarms, and the antennariae and pretentorinae are located more dorsad,| 


markedly so in the former. On the ventral surface, 


there is consid- 


a) 
vt 
oe le 
F fa a 
a VEL. T 
ac 
1G ‘ 
‘ 
ie a 


, Pane > “i Ts ro ; cbr ie a jk The jolfteeome orth at 2B 


m ont LP 
Do, F 
Sait & 
+ ¥ 
| ft 
TE 
~ 
a Tare 
=~ - 
- ‘ 


. 288304 


ab tenos 


toon ead? (fe to tceqte Lent aes edt hating 


a a 
tne ae ony 
—ofee Peas it Loe DP tay: eae one“ aeodth Ame 


ELS abe + ylthetveqde edt ve enageg ib 


afa-are LBAtoes bas _piateinos&) ree vecaconne 


ae —— — a aan i 


mer oyan Sa, Wat aehise debs! a qertad ene ine 
fee rob @d¢ /o1 dieent cinaostan tide 
bithglom s . smmreanirnytids mez LE 


auvne -exeteh eof Ho rod ota om! tokemeee 


} mpitatoast: sit Awe hinis ei oe nibate bode " ‘ 


re miekeyry, y  eeeh bak HLeeSeVaL my Pt Lar Pe tly pers aared 


wens sit 4 Lise, een aes he. nokta 
Sty wh ALL8 Sue. DE ana t* ctnetateat ost all 


" 5%, ok iy Lagvtaset bo es igs 246 Gj =i » ont. senatalitys 


ebiellebroe ost sovetsney dowd Eo oreg 
Prk to suritodustogqdds Aaa Jatat eral ae 
eiitoat ,seveROd 

‘ ptenc unobtet SAF? te ousteants gaT +n 

Ko Belted ashtodsiapsee. odd hy sorttotnanonaeh | 
ant ,eutglzsane nt atelaads S18, Sarre,” 


~ 


1) ee ve. en Pieper 2) sg 9 hat ta Boat cys ee ae 


-~oo S@agooy too of seer earia Gas wats 
ston batevot otis sani tolnetarg ines ba 
at oxen _geetiare Lowney eft: 2s) 


a re ——— ny 
is > _ 


ia 7 rl 7 : . in » - Py , : 


60 


erable variation in the position of the nba cuit arisen The tances | 
ium also varies a great deal. This is probably not a very homogen- ! 
eous group. Sphindus seems to be more structurally similar to Sphae 
sia than any other genus studied. 


Scarabaeoidea.-- This is a very homogeneous group, characterizef 


by a heavy chitinization of the head-wall, degeneration ot the epi- 
eranial suture, ventral migration of the antennariae and the preten- 
torinae, and a complete gula, except in Pseudolucanus, in which the 

| gular sutures extend no more than half the distance between the oc- 
|cipital foramen and the submentum. In the development of the ven- 
tral surface and perhaps the tentorium, the Scarabeeoidea should be 
classed with the Adephaga and related groups. 
Cerambycoidea.-- This is also s very homogeneous group, charac- 
terized by a weak chitinization of the head wall, a distinct dorsal 
position of the antennariae and the pretentorinae, and a more or les 


well-developed epicranial suture. The gular sutures sre generslly 


|short. The tentorium is delicate and membranous throughout. 
Rhynchophora.-- The members of this group mey or may not pboss- 


ess a snout. They may or may not possess confluent gular sutures. 


When they do, the condition is the same as in other Coleoptera. They] 


may or may not possess a labrum. There is no definite character of 


|the head-capsule by which the shynchophora can be separated from 


Jother Coleoptera. The affinities of this group are not at all clear| 


Conclusions.-- The condition of the dorsal surface throughout 


|the groups is most variable, and can be relied upon but little to 


|furnish evidence of the degree of relationship. On the ventral sur-] 


|face the condition is much more stable, and probably much more reli-| 


able in indicéting aifinities. The tentorium is less variable than 


Kot ttl ee ed conttaok “sda 


| 
= yoo Lew a Leeda et ast, beeb, tae 
rus rh od’ tee + EN spo ao Py basen, 
. | | A ia a 
ia cr ERA |; is : 19 Sy oeciien Quio i Utey Bas aki 4 ; aap) 
bi.) it be mio as mre Liaw-peed eat to no nbae ; 
be ‘iene det be has tegnntme any to Hotta cate a 
| bite ; wt anwaoufobsrest ai ¢qecoxe ala ‘Ot a Liane 
fen sty | lath edd then sant - ton on suet 
bith: 6, Pmenge Lovs Pol wo seae es eaiih “hone 
oh twvoxte eehior ada fou gad rvocts ot .-om? aqedisg be 
| égoony befor bie eaedqolh 
netadte kk é-en eatiders un rey cate al of ata 
| - tad fon tt i tt tee ed: at toes ites Lite 
bike atts " axttofass a ant es css xnanetale 
eich Lotus, tating, ofl - pT Ue Letastokee 
ccd idaetdotincbae) siaolias ak iio 
a ta ee ee . am. Oo Hidi peo axxo taba, eat -- 0's 
ranid firs tHeulieos & mod tot yan to * sah Yadh ys 
anietcoel | 7 imeo ont of nold than adt 
4f eced? , eT CRE i s0sseoqy 
e479 Hoatagwis a » etodyontomyal snd 0 Law qe of _ 


te fy ; or ii? to eetitor ta sie i 


faonos sft To amon thneg ai? +o. mee 


¢ sege Aeliog @8 feo, ban,  sisalsay tia nh 
ss : fiaur at? 6 «A ee otal ‘>; ? serge aif? 0, ene! 


:_ 91 (om KAM At fdeduta baa ,eldute es OR sons at sepern 


rad? ofterwmy cent at owlzaliok aff none Led wt ee 
th 


cy ett 


61 
the dorsal surface, and less stable than the ventral surface. Weigh-| 
ine the evidence presented in this study, two large groups of most | 
of the families can be made. The Cupesoidea, the Mordelloidea, the 
EHlateroidea, the Dryopoidea, the Dascilloidea, the Byrrhoidea, the 
Cucujoidea, the Tenebrionoidea, and the Bostrichoidea should probab- 
ly be grouped together, while the charscteristic families of the 
Cantharoidea, the Staphylinoidea, snd the Silphoidea should perhaps 
be grouped with the Adephagea and Hydrophiloidea. The Scsrabaeoidea 
may also be included in this latter group. The affinities of the 
ime eaheeWides and the Khynchophora are too vague to include either 
in the two larger groups. . 

VI. SUMMARY 

1. This investigation deals with the homology of all the struc- 
tures of the head-capsule of one or more representatives of ninety- 
four of the one hundred and nine families of Coleoptere listed by 
Leng in his recent catalogue. One hundred and fifty-five genera, of]}} 
jwhich one hundred and thirty have been figured, nearly all represent} 
ing different important subgroups, have been studied. 

2. This serial study has made it possible to identify the seme 
structures in a wide series of forms, and to definitely fix the ho- 
|jmology of all the parts of the heed-capsule. 

3. Hypothetical types have been constructed, based on the strucy 
liare of the head-capsule of generalized insects and Coleoptera. These 
ishow the Coleoptera to have developed in general a consolidation of 
Isclerites and ea heavier chitinization of the wall of the head, a coms 
|pacting of the heed as a whole, and an approximation of the dorsal | 
jand ventral movable parts at the cephalic end. 


4, The epicrenial suture has been identified in all but two 


¥ ons fer Michal: wand ye cinta: a ds ' oLeide is 


——— 


Sita acer Ca! 


aa? %a Serthimet shfelwetoarade ond oftdw 9, 2ene's 
ener fi ma ih ; if ae } quali la eA? iS | , Pe b l ont redquviate 
my , ne! § OA a “* . a 14 j a | S os ; } f) “7 é sf 5-5 £9 fit _szassaoti sah : 


aS SRA TIES 
=3 ~ 


2 


Eo - oth cn igs 


ytuniA. tal Ravit ete: aay swot TO 6a6” i ‘tubadaiie 


Te mre a ore! pub stds" ab bs 


gat 695 Po Ptabwen ait! Apt onead od? stale ¢ 


t- pot ieaeee ache webees Elona one 20bhd 
vey irnoge soricderiingll oct hae said 


; ~t Wt, ra ae ‘3 £ " 2 y o ] a buh i a “8 ivid Aa be ay 
eronqod oan atl te ; 


+ 
& 
‘ 


. ane eno 
ASUS iv eet Aly _ we 


ide Tce aaa ‘pic oak iw a leen*, cole 


teil date? woeied tec ehitewt: omka bie he sil 
Thos ere eet i ye SeTthaed gag” oopgoLatne een 
he 242 feod eve cities ee 
eed. sree squvrgdae foossode 
sie eee ash2 oF faa soy 2% Sham ane. yout leltew 
ou Tahinitet, 4¢ ore Jeavet to eataea ehiw 
Jo fcumun~t of ade 46 al tay 
qaacd ,bevoratedos aeaeg ovat ene bap tsetd 
Py Hey eye | tou o*nadel ber bre tocen ta: ee > 
t Palace on [wrens ted neqolered Vash: of. 
“ey i acd So {Lee ef? Le doideninleiadg ene 
mos 6 poltentxe waa ae tne (eloty a ee ‘aise a 3 
baw obiodges olf t% biti fa ; 
ow? tue Cia ef Colt tego nate sat Ae a ' 


. 
ia 
‘Tt 7 P 


WY 6 


Li 


of the genera studied. It has proved a great aid in determining the] 
limits of neighboring parts. What may appear to be a distinct epi- | 
cranial suture may not even be a suture. It is sometimes distinctly 
invaginated. Its identity can only be definitely fixed by determin- 
ing the location of the pretentorinae, which are always associated 
with it. 

5. The limits of the vertex are dependent upon the position of 
the epicrenial Sitnieeias In the Khynchophora nearly all of the snout 
belonss to the vertex. 
| 6. The unmodified occipital suture has been identified only in 
the Adephaga. The cephalic end of it on the ventral surface is al- 
jways represented by part of a curving ridge, which is present in all 
but a few genera. 

7. The supratentorinse have peen identified in a few senera, 
nearly all of which belong to the Staphylinoidea. 

8. The pretentorinae are the great landmarks of the head-capsul¢ 
Jand have been identified in all but two genera. They are, in the 
vast majority of genera, located near the cephalic end of the epi- 
eranial arms. A definite determination of the pretentorinae cannot 
always be made without an ental examination of the head, 

9. The size and form of the front is dependent upon the posi- 
ition of the epicranial arms. In the Cerambycoidea it is lerge. In 
many genera, as illustrated by Omophron, Harpalus, and Tachinus, it 
jis partly or wholly invaginated. It may probably be rudimentary of 
|wholly lost in many genere in which the mesal parts of the epicrania} 
aie have disappeared. 


10. What has been celled the clypeal suture in such genera as 


Cicindela and Harpalus is not even a suture, but the line of invagi-| 


a NS tl a a, Ra ee Sa 


aS a 


| Fire c 


ene anaemia 


art i sn 


———————— 


aa 


gies rot 


fsw0e 


se 
vi 
» 4 
} all 
Bi 
" 
a 
i 
‘ 
| 
2 
4 
i 
7 
i] 
et 
\ 
- ie 


RAL! @ 


—— 


bi la | & 


6 ontrage Y of bi ‘sean went” rad oie 
alpen rm *o bp wremt vay te et 


a 


Heit 1 ai 


eat F ue 
i f i a 7 a ii 
hae Oe 
wil Pith 7 “ah 7 ; : 
io an Gog) ae 
: | : ' b 
a i 
if sate 


a 


ry Mnahnogeb co tens Cae 
vats J t Lainie Pics: 
b te tee ot kawighe a 
blew pivris pf te eg xO 
j tee! - 4 AS, 
ae ! (eee ovat) eon RaeRm } 
bud 9 OW snotnd 
" ety att sim ssosrodinut 
ip Gwe red Lis we ‘botgdae 
tase betnser ‘casual : 
¢ ‘tr ty car" ered of nthe fa: 
aninars [ating os toot dw 
10 6! raorvt. «a4 a iret bag 
ednatel ‘etd of .entte fel 
viegiea ,cordgea® yd betatvamiam ve § 
(dedory iam #1 “tetaqigsvnh tulad 
oq. {xGom ec dohse fil sapiihce > 
nae = i ; to 
utc Leegyte ely cna aah: 
oad ,rruteee neve t¢ ) 


rw, 


nation of the front. 


11. The clypeus is always divided into the postclypeus and the 


preelypeus. 


The preclypeus is with one exception always distinctly 
membranous. It may be as large or larger than the labrum. 


12. There id a distinct clypealia present in the Coleoptera, 


and in widely separated groups, such ac the Adephega and the Ceram- 
bycoidea. 
13. The labrum may be indistinctly determined in both Rhyncho- 


|phora and Other Coleoptera. It may also be quite distinct in some 


|Rhynchophorsa where it is considered to be absent. 


14. The submentum is always located distinctly cephaled of the 


occipital foramen, with a chitinized area between it and the forsmen 
15. The metatentorinae may be located on the cephalo-lateral 
border of the occipital foramen, as in generalized insects, or they 
may be far cephalad of this locetion. 
16. All that region between the occipital foramen and the sub- 
mentum is a part of the postgenae, produced by the fusion on the me-]| 


json of the mesal margins of the postgenae. 


17. The gular sutures result fron the cephalic: migration of the} 


metatentorinse. 


18. The gula is that area included between the gular sutures, 
jand is, therefore, derived from the postgenae. The majority of the 
|Coleoptera possess a gula that extends no more than half the distanced 
Joetween the occipital foremen and the submentum. 


| 19. The tentorium of the Coleoptera is typically quite similar 
fin form and development to that of generalized insects. Frequent 


nodifications are loss of chitinization, loss of corpotentorium, and 


Haminatentorium. Occasionelly the pretentorium may be rudimentary. 


és 


. | P ee. S +e hor { L¢ fi ’ ' ‘ é a all syookat 
t 
‘ 


ig it? & reg iY, i te Oy aie: otal heh iv rh: ay aeks 


* 
z = ow 
i ’ 7 , 
a A _ 9 i ‘ as =e bat a 
- LF 


“—_ a . ¥ ere yy 7 _ : if erty Y % 
: } ‘~*~ a ie ¢) s i ei! w oo _" 7 23) ‘ 


a ion i 6 
* y Sy "s a - or 
v 
—- 7 toa 
e a ‘ » r *@ ‘ 
{ a rd P ‘ 9 


Ci j F + f ivaw f i pene Tstv2 one T. Lv el ateqgyle 
: ; - 


TH i “hz Sealieé ty, 3 <. 36 cw ae A 7. 


; lt) 
O's alia } nd act: ! wae J i Te 1G ae 


inet ae J aad Won2dk 6 ee yaar ee: 


ed here iende at thee 

ay i*e toeol ayant Bi. sick 
edt pane J ' » Pazintride = pt Lai es 
ee i jiof’ o¢ Yam ‘aantvoda at h 


= we 
hie sd) — b+ Dexs) ei Sauect Let toes 


i ; dail 
‘ broo0b ainv 2 nat : 


1 


ft gnsenrted noize Fei 


lvotam os .sanentigs ais. ort pov ticwk ’ 

ethan ; 0 tod aig Ae vosewog 

+s? be Otel Logtateve 

bod fas/ “,eFqrdlol ott Ke mini orine? * 
ineavest .ateéani Bociiateties Te Tee taoeqe tome: 
cuisotaedteton io weet ae itvaalel?iie 20 be aol 


- sevavcaniiaos td Gea msi bgt aol og Ae chtane aad 


ee ee oz : 


64 
The functions of the absent parts are assumed by other parts of the 
tentorium, or by the pharynx, or the head may be so compact and chi- 
tinized that a tentorium is no longer needed. 

20. The cephalic migration of the submentum, and the subsequent 
formation of an indistinguishable area between it and the occipital 
foramen is due either to the caudad migration of the occipital for- 
amen, or to the cephalic pull on the mouth-parts, or to both. The 


cephalic migration of the metatentoria, and, therefore, the metaten- 


|torinae, with consequent production of the gula, is probably due to 
| the cephalic pull on the tentorium to furnish a firmer support for 


|the muscles and tendons of the mouth-parts. 


a.4? 


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65 


VI. BIBLIOGRAPHY 
BERLESE, A. 
1909. Gli Insetti. 1: 75-114. Milano. 
BOVING, A., and GHAMPLAIN, A, B. 

1920. Larvae of North American beetles of the family Cleridae.. 

Proc. U.s. Nat. Mus., 57: 575-649; pl. 42-53. 
COMSTOCK, J. H. 

1893. Evolution and taxonomy. Wilder Quarter Century Book. 

PY -St-113; 3 prs. 
COMSTOCK, J. H., and KOCHI, C. 

1902, The skeleton of the head of insects. Amer. Nat., 36: 13- 

45; 29 figs. 
CRAIGHHAD, F. C. 

1920. Biology of some Coleoptere of the families Colydiidae and 
Boethrideridae. Proc. Ent. Soc. Wash., vol. 22, No. 1, 
serpp.}; 2 pis. 

|D'ORCHYMONT, A. 
1916. Notes pour la classification et la phylogenie des Palpi- 


eernia. Ann. Soe, Ent. Fr., 85:° 924106; 6 figs. 


1920. La nervation alaire des Coleopteres. Ann. Soc. Ent. Fr.,] 


89: 1-50; 30 figs; 3 pls. 
GAGE, J. HOWARD. 
1920. The larvae of the Coccinellidae. I11. Biol. Monographs, 
| vol. 6, No. 4., 49 PP; 6 pls. 
GAHAN, C. J. 


| 
| 
! 


] 
} 


1911. On some recent attempts to classify the Coleoptera in ac-| 


cordance with their phylogeny. The Entomologist, 45: 121} 


125, 165-169, 214-219, 245- 248,259-262, 312-314, 348-251, 


| LAMBERE, A, 


| 
| 


66 
592-396; 7 figs. 
GANGLBAUER, L. 
1892- 1904. Die Kafer von Mitteleuropa. Vol. 1-4, Pt. 1. Wien. 
HANDLIRSCH, A. 
1906-1908. Die Fossilen Insekten und die Phylogenie der Rezen- 
ten Formen. P. 1271-80. Leipzig. 
HOPKINS, A. D. 
1915. Preliminary classification of the superfamily Scolytoides 
Tech. Ser., No. 17, Pt. 2., Bur. Hnt., U. S. Dept. Agr., 
Pp. 1654257. 
HYSLOP, J. A. 
1917. The phylogeny of the Hlateridae based on larval charac- 
ters. Ann. Ent. Soc. Amer., vol. 10, No. 3., p. 253-263; 
10 figs. 
KOLBE, H. Jd. 


1901. Vergleichend morphologische Untersuchungen an Coleopteren 


nebst Grundlagen zu einem System und zur Systematik der- | 
selben. Arch. f£. Naturg., p 89-150. 

1908. Mein System der Coleopteren Zeitschr. fur wissenschaft- 
liche Insekten biologie, IV, p. 116-400. 

1911. Die Vergleichende Morphologie und Systematik der Coleop- 
teren, in: Premier Congres international d'Entomologie, 


ii, p. 41-68. 


1900. Notes pour la classification des Coleopteres. Ann. Soc- 


1903. Nouvelles notes pour la classification des Coleopteres. 


Ann. Soc. Ent. Belg. 47: 155-165. 


on 
F 
* 
A 
—_ 


mAPOHeTacd ultweixe eit  ¥o no hteoey Laas: eal sa 
i py soe wv 44 i Py he .* Az < age: ee tHe wt ee 


: 


edu Leviel uiidd sebliefald adi te Gaaaee 


Mes : Ts e% x ath : t ‘ eu _=* “J % e* “one. * ty ag . oat ek . 
s \ 7. = = 


a = 
« ? 
so aa oe 


" atqgefa'l wa ge; o ,dorstetnl edge inqsosqaay nodose 
anxok dts eon? ec: v wadacé aes te ve abse {awe ra 
04{-08 @ , gute > fete .aoe 

es Satyr ww A foe? f fs sat queted ‘tas oer ocr. ake { 


SLD .7 6 elaetold pet ieanl, one eS 
a 


oes ier. *§ . 


S i Sifpastey he sigolotiqiok sbiieissofpagy! 


el =etogotaa' ,LrTudtoeval aertpaoy teineti +s , e178 


am 5 
= * 


~ 


; iL .eotota efed aab aei¢enl iieeert af 1568 netok . 
. s 7 

VeGees Meee 

tqoeiod, 292 esciteoltiegale af Sueg- soton-nel 


6 leiéet rvo Py 8 


2 


67 
LE CONTE, J. L., and HORN, G. H. 
1883. Classification of the Coleoptera of North America. 
Smithsonian Mis. Coll., No. 507. 
LENG, C. W. 
1920. Catalogue of the Coleoptera of America, north of Mexico. 
P, 3-358. Mt. Vernon, N. Y. 
MUIR, F, 
1918. Notes on the ontogeny and morphology of the male genital 
tube in Coleoptera. Trans. Ent. Soc. Lond., 1912: 223-229 
LO pls. 
PETERSON, A. 
1915. Morphological studies on the head and mouth-parts of the 
Biases tore. Ann. Ent. Soc. Amer., 8:.20-67; 7 pls. 
1916. The head-capsule and mouth-parts of Diptera. I11. Biol. 
Mamographs, vol. 3, No. 2, p 1-112; 25 pls. 
RILBY, W. A. 
1904. The embryological development of the skeleton of the head 
of Blatta. Amer. Nat., 38: 777-810; 12 figs. 
SCHIODTE, J. C. 
1861-1883. De lietamorphosi Hleutheratorum Observationes: Bidrag 
til Insekternes Udviklings-historie. 2 vols.; 86 
pls. Kjobenhaven. 
SHARP, D. 
1909. Insects. Cambridge Nat. Hist., 6: 184-298. 
|] SHARP, D., and MUIR, F, 
| 1912. The comparative anatomy of the msle genital tube in Co- 


leoptera. Trans. Ent. Soc. Lond., 1912: 477-639; pl. 


42-78, 


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} 


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Mae <B4e)  tinod ie wink .em ct: orabgoetod) ate 


Te 
ca wt DE 7 ab ee ee Wy, + AG careoilige 
¥? Cl. .weerqit ‘to etsigedteni ban er 
o. .O8 08 bow a in 
abtofedts edt te-reempeleved tagkaeroneene 


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a of cr’ 


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—— 
' 


STRAUS-DURKHEIM, H. HE, 


1828. Considerations generales sur l'anatomie compsree des ani-| 


maux articules, auxquelles on a joint l'anatomie descrip- 


tive du Hanneton vulgaire. P. 51-58; 10 pls. 


HXPLANATION OF PLATE I 


Dorsal 
ips = aca La 
2 


aspect of the head 
Hypothetical head. 
Tetracha carolina. 
Cicindela formosa. 
Calosoma calidum.. 
Omophron americanum. 
Harpalus erraticus. 
Peltodytes 12-punctatus. 
Cybister fimbriolatus. 


Dineutes assSimilis. 


Molamba lunsta. 


Hydrous triangularis. 
Hydrophilus obtusatus. 
Necrophorus carolina. 
Creophilus villo sus. 
Aleochara lata. 
Connophron fossiger. 
Tachinus fimbriatus. 
Leptinus testaceus. 
Stenus flavicornis. 


Gastrolobium bicolor. 


ai 


EXPLANATION OF PLATE ITI 
Dorsal aspect of the head 
Fig. 21. Limulodes psradoxus, 

ee. Sphaerius politus. 

23. Sceaphidium quadriguttatum. 

24. Hister merdarias. 

25. Nacerda melanure. 
Macrosiagon dimidistum. 
Calopteron terminale. 
Photinus pyralis. 
Phengodes plumosa. 
Chaulioenathus pennsylvanicus. 
Collops nigriceps. 
Trichodes nuttali. 
Necrobia rufipes. 

Kpicauta marginate. 
Pytho americanus. 


Cupes concolor. 


Cephaloon lepturides. 


Tomoxia bidenteta. 
Hurystethus debilis. 


Neopyrochroa flabellata. 


PLATS II 


EXPLANATION OF PLATE III 
Dorsal aspect of the head 
Fig. 41. Macratria murine. 
42. Notoxus anchora. 
43. Sandalus niger. 


44. Alaus oculatus. 


Throseus chevrolati. 


Tharops ruficornis. 
Chalcophora virginiensis,. 
Psephenus lecontei. 
-Helichus striatus. 
Eucinetus morio. 

Cyphon rufieellis. 
Stenelmis sinuata. 
Heterocerus undatus. 
Georyssus californicus. 
Dermestes lardarius. 
Byrrhus americanus. 
Nosodendron unicolor. 
Rhysodes americanus. 
Tenebroides sinuatus. 
Phenolia grossa. 
Glischrochilus quadriguttatus. 


Cucujus clayipes. 


PLATE III 


EXPLANATION OF PLATE IV 


Dorsal aspect of the head 


Big. 169. 
64. 
65. 
66. 
Ol 
68. 


70. 


Phyconomus merinus. 
Languria mozardi. 
Megalodacne fasciata. 
Derodontus maculatus. 
Anchicera ephippiata. 
Byturus unicolor. 
Mycetophagus punctatus. 
Philothermus glabriculus. 
Phymaphora pulchella. 
Phalacrus politus. 
Alobates pennsylvanica. 
Melanophthalma angularis. 
Endomychus biguttatus.. 
Hippodamia convergens. 
Arthromacra aenea. 
Pseudocistela brevis. 
Adalia bi-punctata. 
Tenebrio, moli tor. 

Boros unicolor. 
Hyporphagus sp. 


Penthe obliquata. 


PLATE IV 


a a: re ee Ak on oe 
a j ai a = : 
i) Pe eee sh yams er f 
* if i oe ial Z () Pe’ a ie 
Ve a he. 
' p i U 1 LEM 
hd ee er Oe I a 
WE wate’ 
€ 
i tah. €¥ 
“Ae need . 
: * } as 
i 7 ; 


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< 


“THE RRA 
OF THE | | 
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vei q i (ae > 


EXPLANATION OF PLATE V 


Dorsal aspect of the head 


Fig. 84. 
85. 
86. 
87. 
88. 
89. 
90. 
91. 


Ptinus brunneus. 
Sitodrepa panitea. 
Bostrichus bicornis. 
Lyctus planicollis. 
Plesiocis cribrum. 
Sphindus americanus. 
Aphodius fimentarius. 
Dichelonyx elongata. 
Pelidnota punctata. 
Strategus julianus. 
Harpalus erraticus, 
longitudinal cross-section. 
Osmoderma eremicola. 
Trox suberosus. 
Pseudolucanus dama. 
Passalus cornuta. 
Syneta ferruginea. 
Derobrachus brunneus. 
Glycobius speciosus. 
Tetraopes tetraophthalmus. 
Donacia piscatrix. 


Criocerus asparagi. 


EXPLANATION OF PLATE VI 
Dorsal aspect of the head 

Fig..105..Parandra brunnea. 

106. Cryptocephalus quadruplex. 

107. Chrysochus auratus. 

108. Leptinotarsa decimlineata,. 

109. BKupsalis minuta. 

110. Blepharida rhois. 

111. Anoplitis nervosa. 

112. Chelymorpha argus. 


113. Pachymerus arthriticus. 


114. Diabrotica 12-punctata. 


115. Hurymyecter fasciatus. 
116. spicaerus imbricatus. 
117. Rhynehites bicolor. 

118, Dendroctonus valens. 
119. Thecesternus humeralis. 
120. Attelabus analis. 

121. Sphenophorus sequalis 
122. Scolytus quadrispinosus. 


123. Lixus macer. 


EXPLANATION OF PLATE VII 
Lateral aspect of the heed 

Fig. 124, Tetracha carolina. 
125. Cicindela formosa. 
126. Calosoma calidum. 
127. Omophron americanum. 
128. Peltodytes 12-punctatus. 
129. Harpalus erraticus. 
130. Cybister fimbriolatus. 
131. Hydrous triangularis. 


Hydrophilus obtusstus. 


Leptinus testaceus. 


Necrophorus carolina. 
Dineutes assimilis. | 
‘Connophron fossiger. 
Molamba lunata. 
»wtenus flavicornis. 
Gastrolobium bicolor. 
Creophilus villosus. 
Tachinus fimbriatus. 
Aleochara lata. 


Limulodes paradoxus. 


EXPLANATION OF PLATH VIII 
Lateral aspect of the head 
Fig. 144. Sphaerius politus. 
145. Scaphidium quadriguttatum. 
146. Chauliognathus pennsylvanicus, 
cross-section showing invagination 
of part. of. the gula. 
Hister merdarius. 
Photinus pyralis. 
Celopteron terminale. 
Phengodes plumosa. 
Chauliognathus pennsylvanicus, 
Collops nigriceps. 


Trichodes nuttali. 


Necrobia rufipes. 


Cupes concolor. 
Cephaloon lepturides. 
Nacerda melanura. 
Tomoxia bidentata. 
Macrosiagon dimidiatum. 
Epicauta marginata. 
.Kurystethus debilis. 
Neopyrochroa flabellata. 
Pytho americanus, 
Harpalus erraticus, 
dorsal aspect, Showing invagination 


associated with epicranial suture. 


PLATE VIII 


EXPLANATION OF PLATE IX 
Lateral aspect of the head 

Fig. 165. Macratria murina. 

166. Notoxus anchora. 

167. Sandalus niger. 

168. Throsecus chevrolati. 

169. Alaus oculatus. 

170. Tharops rutieerni:s:. 

py fa Heterocerus undatus. 

172. Chalcophore virginiensis, 

173. Georyssus californicus. 

174. Psephenus lecontei. 

175. Helichus striatus. 

Gi Stenelmis Sinuata. 

177, Eucinetus morio. 

178. Gyphon rufiepilis, 

179. Dermestes lardarius. 

180. Byrrhus americanus. 


181. Nosodendron unicolor. 


182. Rhysodes americanus. 


183. Tenebroides sinuatus. 
184. Phenolia grossa. 


185. Glischrochilus quadriguttatus. 


IX 


PLATE 


EXPLANATION OF PLATE X 
Lateral aspect of the head 
Fig. 186. Phyconomus marinus. 


187, Cucujus clavipes. 


188. Languria mozardi. 


189. Megalodacne fasciata. 
190. Anchicera ephippiata. 
191. Byturus unieolor. 
192. Mycetophagus punctatus. 
193, Derodontus asculatus. 
194. Philothermus glabriculus. 
195. Melanophthalma angularis. 
196. Phymaphora pulchella. 
197. Endomychus biguttatus. 
Phalacrus politus. 
Hippodamia convergens. 
Adalia bi-punctata. 
Pseudocistela brevis. 
Alobates pennsylvanica. 
Tenebrio molitor. 
Boros unicolor. 


Arthromacra aenea,. 


EXPLANATION OF PLATE XI 
Lateral aspect of the head 
Fig. 206. Hyporphagus sp. 
207. Penthe obliquata,. 
208. Ptinus brunneus. 
209. Sitodrepa panicea. 
210. Lyectus planicollis. 
2ll. Bostrichus bicornis. 
212. Sphindus americanus. 
213. Plesiocis cribrum. 


214. Aphodius fimetarius. 


215. Dichelonyx elongata. 


216. Pelidnota punctata. 
217. Strategus suiveauee 
218. Osmoderma eremicola. 
219. Trox suberosus. 

220. Pseudolucanus dama. 
221. Passalus cornuta. 
222. Parandra brunnea. 
223. Derobrachus brunneus. 
224. Glycobius speciosus. 


225. Tetraopes tetraophthalmus. 


PLATE XT 


EXPLANATION OF PLATE XII 
Lateral aspect of the head 
Fig. 226. Donacia piscatrix. 
227. Syneta ferruginea. 
228. Criocerus asparagi. 
229. Cryptocephalus quadruplex. 
230. Chrysochus auratus. 
251. Leptinotarsus decimlineata. 
Diabrotica 12-punctatea. 
Blepharida rhois. 


Anoplitis nervosa. 


Chelymorpha argus. 


Pachymerus arthriticus. 
Kupsalis minute. 
Kurymycter fasciatus. 
Rhynchites bicolor. 
Attelabus analis. 
Epicaerus imbricatus. 
Lixus macer. 


243. Thecesternus humeralis. 


EXPLANATION OF PLATE XIII 


Lateral aspect of the head 


Fig. 244. Sphenophorus sequalis. 


245, 


ecolytus quadrispinosus. 


246. Dendroctonus valens. 


Ventral aspect of the head 


Fig. 247. 
248. 
249, 
250. 
251. 
252. 
253. 
254, 
255. 
256. 
257. 
258. 
259. 
260. 
261. 


Hypothetical head. 
Tetracha carolina. 
Cicindela formosa. 
Calosoma calidum. 
Harpalus erraticus. 
Omophron americanum. 
Peltodytes 12-punctatus. 
Cybister fimbriolatus. 
Dineutes assimilis. 
Hydrous triangularis. 
Hydrophilus obtusatus. 
Leptinus testaceus. 
Necrophorus carolina. 
Connophron fossiger. 


Molamba lunata. 


PLATE XIII 


EXPLANATION OF PLATE XIV 
Ventral acpect of the head 
Fig. 262. Stenus flavicornis. 
263. Gastrolobium bicolor. 
264. Creophilus villosus. 
Tachinus fimbriatus. 
Aleochara lata. 
Limulodes paradoxus,. 
Sphaerius politus. 
Scaphidium quadriguttatum. 


Hister merdsarius. 


Calopteron terminale. 


Photinus pyralis. , 

Phengodes plumosa. 
Chauliognathus pennsylvanicus. 
Cantharis bilineatus,. 

Collops nigriceps. 

Trichodes nuttali. . 

Necrobia rufipes. 


Cupes concolor. 


PLATE XIV 


EXPLANATION OF PLATE XV 


Ventral aspect of the head 


Fig. 280. 
281. 
282. 
283. 
284. 
285. 
286. 
287. 
288. 
289. 
290. 
201. 
292. 
293. 
294, 
295. 
296: 


Cephaloon lepturides. 
Nacerda melanura. 
Tomoxia bidentata. 
Macrosiagon dimidtatum,. 
Kpicauta marginata. 
Kurystethus debilis. 
Pytho americanus. 
Neopyrochros flabellata. 


Macratria murina. 


Sandalus niger. 


Notoxus anchora. 

Alaus oculatus. 

Thearops ruficornis. 
Throscus chevrolati. 
Chalcophora virginiensis. 
Psephenus lecontei.. 


Helichus striatus. 


PLATE XV 


vy: 


EXPLANATION OF PLATE XVI 


Ventral aspect of the head 


ie. 297. 


298. 
rk 
300. 
301. 
302. 
503. 
304. 
505. 
506. 
307. 
308. 
509. 
310. 
or, 
312. 
513. 
314. 
$15. 


Stenelmis sinuata. 
Heterocerus undatus. 
Georyssus californicus. 
Encinetus morio. 

Cyphon ruficollis. 
Dermestes lardarius. 
Byrrhus americanis. 
Nosodendron unicolor. 
Rhysodes americanum. 
Tenebroides sinuatus. 
Phenolia grossa, 
Glischrochilus quadriguttatus. 
Phyconomus marinus. 
Cucujus clavipes. 
Languria mozeardi. 
Megalodacne fasciata. 
Derodontus maculatus. 
Anchicera ephippiata. 


Byturus unicolor. 


yier site 


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EXPLANATION OF PLATE XVII 
Ventral aspect of the head. 

Fig. 316. Mycetophagus punctatus. 

317. Philothermus glabriculus, 

318. Melanophthalma angularis. 

319. Phymaphora pulchella. 

320. Endomychus biguttatus. 

$21. Phalacrus politus. 

322. Hippodamia convergens. 

323. Adalia bi-punctata. 

324. Pseudocistela brevis. 

525. Alobates pennsylvanicea. 

426. Tenebrio molitor. 

427. Boros unicolor. 

328. Arthromacra aenea, 

429. Hyporphagus sp. 

330. Penthe obliquata. 

551. Ptinus brunneus. 

332. Sitodrepa panicea. 


433. Bostrichus bicornis. 


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PLATE XVII 


EXPLANATION OF PLATE XVIII 
Ventral aspect of the head 
Fig. 334. Lyetus planicollis. 
455. Sphindus americanus. 
456. Plesiocis cribium. 
357. Aphodius fimetarius. 
338. Dichelonyx elongata. 
339. Pelidnota punctata. 
440. Strategus julianus. 


441. Osmoderma eremicola. 


$42. Trox suberosus. 


343. Pseudolucanus dama. 

444, Passalus cornuta. 

345. Parandra brunnea. 

246. Derobrachus brunneus. 

447. Glycobiws speciosus. 

548. Tetraopes tetraophthalmus, 
349. Donacia piscatrix. - 

450. Syneta ferroginea. 

451. Criocerus asparagi. 


352. Cryptocephalus quadruplex. 


is 
on 
HH 
> 
Sq 
2 
i= 
=< 
a 
yy 


EXPLANATION OF PLATE XIX 
Ventral aspect of the head 
Fig. 353. Chrysochus auratus. 


454. Leptinotarsa decimlineata. 


355. Diabrotica 12-punctata. 


556. Blepharida rhois. 

457. Anoplitis nervosa. 

358. Chelymorpha argus. 

359. Pachymerus arthriticus. 
560. Eupsalis minuta. 

561. Kurymycter Facelavuee 
562, Rigneiites bieoLlor. 

. Attelabus analis. 
EKpicserus imbricatus. 
eecolytus quadrispinosus. 
Lixus macer, 
Thecesternus humeralis. 
Sphenophorus aequalis. 


Dendroectonus valens. 


(statssn 


EXPLANATION OF PLATE XX 
Endoskeleton of the head 
Fig. 370. Hypothetical head. 
371. Tetracha carolina. 
372. Cicindela formosa. 
Calosoma calidum. 
Harpalus erraticus. 
Omophron americanum, 
Peltodytes 12-punctata. 


Cybister fimbriolatus. 


Dineutes assimilis. 


Hydrous triangulsaris. 
Hydrophilus obtusatus. 
Leptinus testaceus. 
Necrophorus carolina. 
Connophron fossiger. 
Stenus flavicornis. 
Gastrolobium bicolor. 


Creophilus villosus. 


EXPLANATION OF PLATE XXI 


Endoskeleton of the head 


Fig. 


387. 
588. 
389. 
590. 
S91. 
392. 
393. 
594. 
3595. 
596. 
597. 


398.: 


399. 
400. 
401. 
402. 
403. 


Tachinus fimbriatus. 
Aleochara lata, 

Limuiodes paradoxus. 
Sphaerius politus. 
Scaphidium quadriguttatum. 
Hister merdarius. 
Calopteron terminale. 
Photinus pyralis. 
Phengodes plumosa. 

Ciesla opus thud pennsylvanicus, 
Collops nigriceps. 
Trichodes nuttali. 
Necrobia rufipes. 

Cupes concolor. 

Cephaloon lepturides,. 
Nacerda melanura, 


Tomoxia bidentata. 


PLATE 


XXI 


EXPLANATION OF PLATE XXII 

Endoskeleton of the heed. 

Fig. 404. Macrosiagon dimidiatum. 

405. Epicauta marginata. 
406. Eurystethus debilis. 
407. Pytho americanus. 
408. Neopyrochroa flabellata. 
409. Macratriea murina. 
410. Notoxus anchora. 
411. Sandalus niger. 


412. Alaus oculatus. 


413. Tharops ruficornis. 


414, Throscus chevrolati. 

415. Chalcophora virginiensis. 
416. Psephenus lecontei. 

417. Helichus striatus. 


418. Stenelmis sinuata. 


a 


EXPLANATION OF PLATH XXIII 
Hndoskeleton of the head 
Fig. 419. Heterocerus undatus. 
420. Georyssus californicus. 
421. Eucinetus morio. 
Cyphon ruficollis. 
Dermestes lardarius. 
Byrrhus americanus. 
Nosodendron unicolor. 


Rhysodes americanus, 


Tenebroides sinuata. 


Phenolia grossa. 
Glischrochilus quadriguttatus. 
Phyconomus marinus. 

Cucujus clavipes. 

Languria mozardi. 

Megalodacne fasciatus. 
Derodontus maculatus. 


Anchicera ephippiata. 


PLATE XXIII 


EXPLANATION OF PLATE XXIV 
EKndoskeleton of the head 
Fig. 436. Byturus unicolor. 
457. Mycetophagus punctatus. 
438. Philothermus glabriculus. 
439. Melanophthalma angularis. 


440. Phymaphora pulchella. 


441. Endomychus biguttatus. 


442, Phalacrus politus. 

443. Hippodamia convergens. 
444, Adalia bi-punctata. 
445, Pseudocistela brevis. 
446. Alobates pennsylvanica. 
447, Tenebrio molitor. 

448. Boros unicolor. 

449, Arthromacra eenea, 

450. Hyporphagus sp. 

451. Penthe obliquata. 


EXPLANATION OF PLATE ZXXV 
Endoskeleton of the head 
Fig. 452. Ptinus brunneus. 
453. Sitodrepa panices. 
454. Bostrichus bicornis. 
455. Lyctus planicollis. 
456. Sphindus smericanus. 
Plesiocis cribrum. 
Aphodius fimetarius. 


Dichelonyx elongata. 


Pelidnota punctata. 


Strategus julianus. 
Osmoderma erimicola. 
Trox suberosus. 
Pseudolucanus dama. 
Passalus cornutus. 
Passalus cornutus, 


lateral aspect, cross-section. 


BXPLANATION OF PLATE XXVI 


Endoskeleton of the head 


Fig. 467. 
468. 
469. 
470. 
All. 
A72. 
473, 
A74, 
A75, 
476. 
A77. 
A478, 
479. 
480. 
481. 


Parandra brunnea. 
Derobrachus brunneus. 
Glycobius sSpeciosus. 

pet enenee tetraophthalmus,. 
Donacia piscatrix. 

Syneta ferruginea. 
Criocerus asparagi. 
Cryptocephalus quadruplex. 
Blepharida rhois. 
Chrysochus auratus. 
Leptinus testaceus. 
Diabrotica 12-punctata. 
Anoplitis nervosa. 
Pachymerus arthriticus. 


Chelymorpha argus. 


PLATE XXVI 


EXPLANATION OF PLATE. XXVITI 
Endoskeleton of the head 
Fig. 482. Kupsalis minuta. 
483. Kurymycter fasciatus. 
484.xhynchites bicolor. 
485. Attelabus analis. 


486. Kpicaerus imbricatus. 


487. Thecesternus humeralis. 


488. Lixus macer. 
489. Sphenophorus aequalis. 
490. Scolytus quadrispinosus. 


491. Dendroctonus valens,. 


a 


Segre 
= 
= “ 
Las 
= 
re 
= 


VITA 

The author of this thesis, Fenner Satterthwaite Stickney, was 
born in Wilson, North Carolina, August 6, 1892. He attended the 
public schools of Wilson, until May 1909. The following year he 
spent on a farm in Beaufort County, North Carolina. During the sum 
mer of 1910, he moved to the Hawaiian Islands, and entered in Sep- 
tember of that year the McKinley High School of Honolulu, from whic 
|} he graduated in June 1912. From September 1912 until June 1914, he 
| Studied agriculture, including a course in Entomology, at the Col- 
lege, now the University, of Hawaii. In August 1914, he registered 
} in the College of Agriculture of the University of California; he 
majored in Entomology, and received the degree of Bachelor of Sci- 


ence in Agriculture from that University in May 1916. The year 1916 


-1917 he spent on a farm in Brunswick County, North Carolina, en- 


gaged in practical agriculture. He attended the Summer Session of 
1917 at the University of Illinois, specializing in economic Ento- 
mology. During the year 1917-18, he was registered in the Graduate | 
school of the University of Illinois, from which he received the de 
gree of Master of Science in tntomology in June 1918. June 1, 1918,} 
he was inducted into the United States Army and ordered to Camp 

1 Dodge, Iowa, where he was engaged in sanitary entomological work un 

| til August 1918, when he was sent oversess. After six months in 

? Frence as a member ot Sanitary Squad #70, he wes returned to the U- : 
| nited States, and received his discharge in Honolulu in June 1919. 

) From September 1919 to June 1921 he wes a Fellow in Entomology at 

| the University of Illinois. During the summer ot 1920 he assisted 

| in extension entomological work at the North Dakota Agricultural 


College, 


Fig.492. 
493, 
494, 
495. 
496. 
497, 
498. 
499, 
500. 
501. 
502. 
503. 


EXPLANATION OF PLATE XXVIII 


Amphizoa 
Amphizoa 
Amphizoa 
Amphizoa 
Pilepius 
Pilopius 
miley ins 
Paidopiws 
Othnius 

Othnius 

Othnius 


Othnius 


lecontei, dorsal aspect. 
lecontei, lateral aspect. 
lecontei, ventral aspect. 
lecontei, endoskeleton. 
lacustris, dorsal aspect. 
lacustris, lateral aspect. 
lacustris, ventral aspect. 
lacustris, endoskeleton. 
sp., dorsal aspect. 

Sp., lateral aspect. 

sp., ventral aspect. 


sp., endoskeleton. 


ey 


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PLATE XXVIII 


